Ecology and regeneration of forest communities on the alluvial fan ...

Ernst Moritz Arndt University Greifswald 

Institute <strong>of</strong> Botany <strong>and</strong> L<strong>and</strong>scape <strong>Ecology</strong> 

Diploma thesis in the study program <strong>of</strong> L<strong>and</strong>scape <strong>Ecology</strong> <strong>and</strong> Nature Conservation 

<strong>Ecology</strong> <strong>and</strong> <strong>regeneration</strong> <strong>of</strong> <strong>forest</strong> <strong>communities</strong> on the 

alluvial fan near Qakh (Northwestern Azerbaijan) 

by 

Michael Zimmermann 

-March 2009- 

Supervised by: 

Pr<strong>of</strong>. Dr. Michael Manthey <strong>and</strong> Pr<strong>of</strong>. Dr. Michael Succow

Hiermit erkläre ich die vorliegende Diplomarbeit mit dem Thema: 

„<strong>Ecology</strong> <strong>and</strong> <strong>regeneration</strong> <strong>of</strong> <strong>forest</strong> <strong>communities</strong> on the alluvial fan near Qakh 

(Northwestern Azerbaijan)“ 

selbstständig verfasst und keine <strong>and</strong>eren Hilfsmittel als die angegebenen verwendet zu 

haben. Aus <strong>and</strong>eren Werken in Wortlaut oder Sinngehalt entnommene Inhalte sind 

durch Quellenverweis, auch für Sekundärliteratur, kenntlich gemacht. 

Greifswald, den 13.3.2009 


Kontakt: 


Grimmer Str. 70 

17489 Greifswald 

zimmermannmicha@gmx.de

Danksagung 

Hiermit möchte ich mich bei all denen bedanken, die zum Gelingen dieser Arbeit 

beigetragen haben. Arne Truckenbrodt und Stefan Israel danke ich für ihre tatkräftige 

Unterstützung bei der Feldarbeit, Sebastian Schmidt für die gute Betreuung vor Ort, 

Hagen Gottschling für die Einführung ins Gebiet, Familie Okradjanashvili für 

Unterkunft und Hilfe vor Ort. 

In fachlicher Hinsicht habe ich vor allem Jan Peper zu danken, der immer mindestens 

ein <strong>of</strong>fenes Ohr für Probleme hatte, desweiteren meinen Betreuern Pr<strong>of</strong>. Dr. Michael 

Manthey und Pr<strong>of</strong>. Dr. Michael Succow. Florian Jansen danke ich v.a. für technische 

Unterstützung aller Art. 

Dank auch an Maria Langhammer, Dorothea Pietsch, Susanne Abel, Frederick Noack 

und Martin Wiehle, die lange Zeit meine Leidens- und Diskussionspartner im 

Diplom<strong>and</strong>enzimmer des Botanischen Instituts waren. 

Ein ganz besonderer Dank geht an meine Eltern, die mich während des ganzen 

Studiums in jeglicher Hinsicht unterstützt haben sowie an meine Familie Annette und 

Moritz, mit der ich nun wieder mehr Zeit verbringen kann. 

Abschließend möchte ich mich bei der Succow-Stiftung und beim DAAD bedanken, 

deren Stipendien mir erst die finanzielle Möglichkeit gaben, diese Arbeit in Angriff zu 

nehmen.

Index 

1 Introduction...............................................................................................................1 

2 Study area...................................................................................................................3 

2.1 Location....................................................................................................................3 

2.2 Relief <strong>and</strong> Geology...................................................................................................4 

2.3 Climate......................................................................................................................5 

2.4 Hydrology.................................................................................................................6 

2.5 Vegetation, soils <strong>and</strong> l<strong>and</strong> use..................................................................................6 

3 Materials <strong>and</strong> Methods.............................................................................................8 

3.1 Vegetation sampling.................................................................................................8 

3.2 Rejuvenation sampling.............................................................................................9 

3.3 St<strong>and</strong> structure .........................................................................................................9 

3.4 Site Parameters.......................................................................................................10 

3.5 Data analysis...........................................................................................................10 

4 Results.......................................................................................................................13 

4.1 Vegetation <strong>communities</strong>.........................................................................................13 

4.2 Rejuvenation...........................................................................................................27 

4.3 Ordination results ..................................................................................................33 

4.4 Ordination results (Successional trends)................................................................35 

5 Discussion.................................................................................................................37 

5.1 Vegetation zonation <strong>and</strong> site conditions.................................................................37 

5.2 Successional trends.................................................................................................45 

5.3 The human impact on the ecosystem......................................................................47 

5.4 Outlook...................................................................................................................51 

6 Summary..................................................................................................................52 

7 Zusammenfassung...................................................................................................54 

8 Literature.................................................................................................................56 

Appendix.........................................................................................................................60

Figure index 

Fig. 1.1 Natural river dynamics on the alluvial fan...........................................................2 

Fig. 2.1 Map <strong>of</strong> alluvial fan...............................................................................................3 

Fig. 2.2 Geological map <strong>of</strong> Qakh region...........................................................................4 

Fig. 2.3 Outline <strong>of</strong> an alluvial fan......................................................................................5 

Fig. 2.4 Climate chart for Zaqatala....................................................................................6 

Fig. 3.1 Location <strong>of</strong> the 71 vegetation plots......................................................................8 

Fig. 3.2 Vegetation sampling design.................................................................................9 

Fig. 3.3 Point Centred Quarter Method (PCQ)................................................................10 

Fig. 4.1 Dendrogram <strong>of</strong> the plant <strong>communities</strong>...............................................................13 

Fig. 4.2 Importance values <strong>of</strong> Equisetum telmateia-Alnus barbata community.............15 

Fig. 4.3 Equisetum telmateia-Alnus barbata <strong>forest</strong>.........................................................16 

Fig. 4.4 Importance values <strong>of</strong> Fraxinus excelsior-Pterocarya pterocarpa community..16 

Fig. 4.5 Fraxinus excelsior-Pterocarya pterocarpa <strong>forest</strong>..............................................17 

Fig. 4.6 Absolute tree density <strong>and</strong> total basal area..........................................................18 

Fig. 4.7 Boxplots.............................................................................................................18 

Fig. 4.8 Importance values <strong>of</strong> Swida sanguinea-Populus alba community....................19 

Fig. 4.9 Populus alba st<strong>and</strong>.............................................................................................20 

Fig. 4.10 Importance values <strong>of</strong> Oplismenus undulatifolius-Acer velutinum community 20 

Fig. 4.11 Acer velutinum <strong>and</strong> Fraxinus excelsior............................................................21 

Fig. 4.12 Importance values <strong>of</strong> Quercus robur-Carpinus betulus community...............22 

Fig. 4.13 Quercus robur-Carpinus betulus community..................................................23 

Fig. 4.14 Importance values <strong>of</strong> Quercus iberica-Carpinus orientalis community.........23 

Fig. 4.15 Multi stemmed Carpinus orientalis.................................................................24 

Fig. 4.16 Importance values <strong>of</strong> Scleropoa rigida-Populus nigra community................25 

Fig. 4.17 Populus nigra st<strong>and</strong>..........................................................................................26 

Fig. 4.18 Rejuvenation sampling.....................................................................................27 

Fig. 4.19 Rejuvenation data 1..........................................................................................30 



Fig. 4.22 Ordination diagram (DCA)..............................................................................33 

Fig. 4.23 Ordination diagram (DCA)..............................................................................34 

Fig. 4.24 Ordination diagram (NMDS)...........................................................................35 

Fig. 5.1 Rough map <strong>of</strong> the alluvial fan............................................................................37

Fig. 5.2 Boxplots.............................................................................................................38 

Fig. 5.3 Longitudial section.............................................................................................39 

Fig. 5.4. Cross section <strong>of</strong> the upper fan ..........................................................................40 

Fig. 5.5 Cross section <strong>of</strong> the lower fan............................................................................41 

Fig. 5.6 Cross section <strong>of</strong> the riverbed (upper fan)...........................................................42 

Fig. 5.7 Cross section <strong>of</strong> the river bed (mid-fan).............................................................43 

Fig. 5.8 Cross section <strong>of</strong> the riverbed (lower fan) ..........................................................44 

Fig. 5.9 Successional development <strong>of</strong> the <strong>forest</strong> <strong>communities</strong> .......................................45 

Fig. 5.10 Flood protection measures...............................................................................47 

Fig. 5.11 Gravel dredging................................................................................................48 

Fig. 5.12 Recently logged Populus alba..........................................................................49 

Fig. 5.13 Wood pasture....................................................................................................50 

Fig. 5.14 Plantations........................................................................................................51 

Fig. 6.1 River dynamics on the upper fan........................................................................53 

Fig. 7.1 Natural riverbed..................................................................................................55

Table index 

Table 3.1 Transformed abundance data...........................................................................11 

Table 3.2 Species included in the NMDS analysis .........................................................12 

Table 4.1 Percentage synoptic table................................................................................14 

Table 4.2 Site <strong>and</strong> structure parameters for all <strong>communities</strong> ........................................15 

Table 4.3 Correlation between selected environmental parameters <strong>and</strong> DCA axes .......33 

Table 4.4 Correlation between selected environmental parameters <strong>and</strong> DCA axes ......34 

Table 5.1 Flooding tolerance classes...............................................................................40

Abbreviations I 

a.s.l above sea level 

°C degree Celsius 

cm centimetre 

com. community 

DBH diameter at breast height 

DCA detrended correspondence analysis 

fig. figure 

g 

gram 

ha hectare 

kg kilogram 

km kilometre 

m 

metre 

mm millimetre 

NMDS non-metric multidimensional scaling 

PCQ point centred quarter method 

spec. species 

spp. species (Pl) 

t 

tonne

Abbreviations II – plant names 

ACE-CAM 

ACE-SPP 

ACE-VEL 

ALN-BAR 

CAR-BET 

CAR-ORI 

CAR-SPP 

COL-ARB 

COR-AVE 

COR-MAS 

COR-SPP 

COT-COG 

COT-SAX 

CRA-SPP 

CYD-OBL 

DIO-LOT 

EUO-EUR 

FIC-CAR 

FRA-EXC 

HIP-RHA 

JUG-REG 

LIG-VUL 

MAL-SPP 

MES-GER 

MOR-SPP 

POP-ALB 

POP-NIG 

PRU-CER 

PTE-PTE 

PYR-COC 

QUE-IBE 

QUE-ROB 

QUE-SPP 

ROB-PSE 

SAL-SPP 

SÄM-SPP 

SOR-TOR 

SWI-SAN 

ULM-FOL 

VIB-OPU 

Acer campestre 

Acer spp. 

Acer velutinum 

Alnus barbata 

Carpinus betulus 

Carpinus orientalis 

Carpinus spp. 

Colutea arborescens 

Corylus avellana 

Cornus mas 

Cornus spp. 

Cotinus coggygria 

Cotoneaster saxatilis 

Crataegus spp. 

Cydonia oblonga 

Diospyros lotus 

Euonymus europaeus 

Ficus carica 

Fraxinus excelsior 

Hippophae rhamnoides 

Juglans regia 

Ligustrum vulgare 

Malus spp. 

Mespilus germanica 

Morus spp. 

Populus alba 

Populus nigra 

Prunus cerasus 

Pterocarya pterocarpa 

Pyracantha coccinea 

Quercus iberica 

Quercus robur 

Quercus spp. 

Robinia pseudoacacia 

Salix spp. 

unknown seedling 

Sorbus torminalis 

Swida sanguinea 

Ulmus foliacea/minor 

Viburnum opulus

-Introduction- 

1 Introduction 

Alluvial <strong>forest</strong>s are considered to be the <strong>communities</strong> with the highest diversity in 

species <strong>and</strong> with the highest degree <strong>of</strong> organisation (Schnitzler 1994). The nature 

conservation value <strong>of</strong> riparian zones <strong>and</strong> alluvial <strong>forest</strong>s has been emphasized during the 

past by several authors (Peterken & Hughes 1995; Gilvear et al. 2000; Hughes & Rood 

2003; Peper 2007). Alluvial <strong>forest</strong>s serve as an interface between two very distinctive 

ecosystems, terrestrial <strong>and</strong> aquatic, across which energy <strong>and</strong> matter (in the form <strong>of</strong> 

nutrients, sediment <strong>and</strong> water) regularly move in both directions. Fluvial 

geomorphological processes are responsible for the development <strong>of</strong> a highly dynamic 

<strong>and</strong> rapidly changing l<strong>and</strong>form surface. This results in a complex mosaic <strong>of</strong> different 

vegetation types with species distributed in response to their tolerances to soil moisture 

<strong>and</strong> nutrient conditions dictated by flood regime <strong>and</strong> sedimentation patterns (Kupfer & 

Malanson 1993). However, the development <strong>of</strong> alluvial <strong>forest</strong>s has been affected by 

human activities for a long time. Throughout the temperate zone alluvial <strong>forest</strong> 

biodiversity has been lost due to damaging activities which include canalization, 

construction <strong>of</strong> flood embankments, agriculture, <strong>and</strong> removal <strong>of</strong> woodl<strong>and</strong> (Petts & 

Moller 1989). In the northwestern part <strong>of</strong> Azerbaijan numerous rivers transport material 

from the adjacent mountain zones <strong>of</strong> the Greater Caucasus <strong>and</strong> build up extensive foot 

slopes along its southern mountain fronts. These foot slopes composed <strong>of</strong> fluvial 

sediments are known as alluvial fans. The natural vegetation <strong>of</strong> these alluvial fans has 

been significantly altered by l<strong>and</strong> management. With the development <strong>of</strong> settlements, 

grassl<strong>and</strong>s, agricultural fields, plantations <strong>and</strong> flood control measures the major part <strong>of</strong> 

woodl<strong>and</strong> areas has been lost or severely altered. Compared to other fans <strong>of</strong> this region 

the quite large alluvial fan near Qakh still provides a relatively high amount <strong>of</strong> alluvial 

<strong>forest</strong>. Due to the fact that the alluvial <strong>forest</strong> <strong>communities</strong> <strong>of</strong> this region are not 

described in detail, yet <strong>and</strong> due to the persistent loss <strong>of</strong> natural woodl<strong>and</strong> as a result <strong>of</strong> 

the above described reasons, the aim <strong>of</strong> this study was to analyse the <strong>forest</strong> <strong>communities</strong> 

on the alluvial fan near Qakh, which are representative for this region. Based on the 

above mentioned facts the following questions were adressed: 

-<strong>Ecology</strong> <strong>and</strong> <strong>regeneration</strong> <strong>of</strong> <strong>forest</strong> <strong>communities</strong> on the alluvial fan near Qakh (Northwestern Azerbaijan)- 

1

-Introduction- 

1. What is the structure <strong>and</strong> species composition <strong>of</strong> the existing <strong>forest</strong> 

<strong>communities</strong> 

2. What are the required site conditions <strong>of</strong> these <strong>communities</strong> 

3. Are the <strong>communities</strong> connected by successional trends 

Fig. 1.1 Natural river dynamics on the alluvial fan with Greater Caucasus in the background 


2

-Study area- 

2 Study area 

2.1 Location 

The study area is situated at 41°22' N <strong>and</strong> 46°51' E on the southern foothills <strong>of</strong> the 

Greater Caucasus <strong>and</strong> is part <strong>of</strong> the Alasan-Awtoran depression (northwestern region <strong>of</strong> 

Azerbaijan). It is an alluvial fan complex, which lies downstream <strong>of</strong> the small town <strong>of</strong> 

Qakh. The altitude within the surveyed area ranges from 177 to 489 metres above sea 

level. The fan consists <strong>of</strong> fluvial sediments, that eroded from the adjacent mountain 

ranges <strong>and</strong> were deposited in an extensive foot slope along the mountain front. The 

sediments are transported by the river Kurmukcay. The study site is surrounded up to 

the north, south east <strong>and</strong> southwest by roads. One road cuts across the mid-fan. Another 

one cuts across the lower fan, as does the railroad line Baku-Zaqatala. Some small 

villages are scattered over the fan, even in its central region. 

N 

4 k m 

Fig. 2.1 Map <strong>of</strong> alluvial fan complex at the southern slope <strong>of</strong> the Eastern Caucasus (Map source: 

(Skworzow 1976)). Inset: Map <strong>of</strong> Azerbaijan <strong>and</strong> location <strong>of</strong> the study area (Map source: (Kurtubadze 

2008)) 


3

-Study area- 

2.2 Relief <strong>and</strong> Geology 

The southern slope <strong>of</strong> the Eastern Caucasus is composed <strong>of</strong> Jurassic <strong>and</strong> Palaeogene 

sediments. Due to the elevation <strong>of</strong> the mountains <strong>and</strong> the steep slopes recent erosion is 

considerable. Hence, large amounts <strong>of</strong> sediments are moved by the rivers. 

Transportation rates increase from the western to the eastern part <strong>of</strong> the mountains, 

reaching maximum values between 2 kg/m³ <strong>and</strong> 5 kg/m³ in the Eastern Caucasus. This 

intensification is due to the the increasing portions <strong>of</strong> slates <strong>and</strong> marls <strong>and</strong> the inferior 

<strong>forest</strong> cover. In addition debris flows are quite common in various parts <strong>of</strong> the Caucasus 

Mountains. In many regions they are regarded as the most powerful process <strong>of</strong> 

denudation. Their appearance is also increasing to the East, because <strong>of</strong> the more 

continental climate, the decreasing <strong>forest</strong> cover <strong>and</strong> the more frequent occurrence <strong>of</strong> 

easy removable Meso- <strong>and</strong> Cenozoic rocks (Franz 1973). 

N 

2 0 k m 

Fig. 2.2 Geological map <strong>of</strong> Qakh region 

(blue: Jurassic, green: Cretaceous, grey dotted: Pleistocene, grey chequered: Pleistocene/Holocene.) (Map 

source: (Alizade 2008)) 


4

-Study area- 

Alluvial fans are typical l<strong>and</strong>forms <strong>of</strong> regions with a considerably high amount <strong>of</strong> 

mechanical weathering. They are composed <strong>of</strong> fluvial sediments build up by means <strong>of</strong> 

river sedimentation processes (Fig. 2.3). They preferentially appear in semiarid areas, at 

the foot slopes <strong>of</strong> high mountain ranges <strong>and</strong> also in periglacial regions. Not the whole 

fan surface is active at a given time. Eventually, accumulation <strong>of</strong> sediments downstream 

makes it easier for a new flood to flow over another part <strong>of</strong> the fan. This switch from 

one part to another is called avulsion. The typical cone shape <strong>of</strong> most alluvial fans 

results from regular avulsion <strong>of</strong> the river. Due to decreasing erosion <strong>and</strong> transportation 

rates <strong>of</strong> the rivers towards their distal margins a characteristic series <strong>of</strong> sediments is 

given. The coarsest material is dropped at the top <strong>of</strong> the fan, s<strong>and</strong> <strong>and</strong> gravel in the midfan, 

<strong>and</strong> fine material at the lower reaches (Ahnert 1996). 

Fig. 2.3 Outline <strong>of</strong> an alluvial fan (Strahler 1975) 

2.3 Climate 

The warm temperate climate <strong>of</strong> the Alasan-Awtoran depression with sufficient rainfall 

during the whole vegetation period provides excellent conditions for <strong>forest</strong> growth. The 

Greater Caucasus mountain ranges protect it from cold weather <strong>and</strong> strong winds. 

Duration <strong>of</strong> the vegetation period varies between eight <strong>and</strong> ten months. Spring, summer 

<strong>and</strong> fall receive each about one third <strong>of</strong> the annual rainfall. Mean temperature in January 

is 0°C, the mean temperatures in July range from 22,5°C to 25°C. The end <strong>of</strong> summer 


5

-Study area<strong>and</strong> 

winter are the driest periods <strong>of</strong> the year. Snowfall rate at the southern slope is 

generally low. Once a decade very cold winters with strong frost periods occur, which 

restrains the growth <strong>and</strong> culture <strong>of</strong> frost intolerant species (Prilipko 1954). 

Fig. 2.4 Climate chart for Zaqatala, 40 km northwest <strong>of</strong> Qakh, with climatic classification after Köppen 

(Mühr 2007) 

2.4 Hydrology 

The 55 km long river Kurmukcay is a w<strong>and</strong>ering gravel bed river with the increasing 

tendency towards braiding at the fan's distal margin. It drains 560 km² <strong>of</strong> the adjacent 

mountain region. Its spring is located in the higher mountain region in more than 

3,000 metres above sea level. Its upper reaches are mainly fed by snow <strong>and</strong> glacier melt 

water, whereas the proportion <strong>of</strong> rainwater increases with decreasing altitude 

(Administration <strong>of</strong> Water Resources Qakh 2007). At the southern slopes <strong>of</strong> the Greater 

Caucasus flood events mainly occur due to snowmelt in springtime. At this time annual 

water run-<strong>of</strong>f reaches its maximum (Franz 1973). Though various measures are taken by 

the government to reduce the constitutive power <strong>of</strong> this river system, active braided 

river channels constantly change the fan's surface. In the warm summer period 

discharge rates strongly decline <strong>and</strong> the riverbed becomes largely dry. 

2.5 Vegetation, soils <strong>and</strong> l<strong>and</strong> use 

According to Meusel et al. (1965) the Alasan-Awtoran depression belongs to the 

southeastern trans-Caucasian province <strong>of</strong> the Caucasian sub region, which is in turn part 


6

-Study area<strong>of</strong> 

the Macaronesic-Mediterranean region. In the past, broad leafed <strong>forest</strong>s constituted 

the natural vegetation <strong>of</strong> this region. Today only small <strong>forest</strong> patches are left. They 

totally cover about one third <strong>of</strong> the study area. They act as an intermediary between the 

European <strong>forest</strong> vegetation <strong>and</strong> the far more species-rich South Caspian Rain<strong>forest</strong>s. 

Heading south they are soon displaced by semiarid steppe vegetation, whereas heading 

north by montane beech <strong>and</strong> lime tree <strong>forest</strong>s. These alluvial <strong>forest</strong> remnants alternate 

with small villages, agriculture fields <strong>of</strong> tobacco, wheat, corn, melon, plantations <strong>of</strong> 

hazelnut, walnut but also mulberry <strong>and</strong> different fruit trees. Cattle, sheep, goat <strong>and</strong> 

water buffalo are the most important livestock species in the fan area. The typical 

spatial pattern <strong>of</strong> different substrates on the fan results in very different silvicultural 

characteristics. The deep loamy alluvial soils at the lower reaches are moist <strong>and</strong> nutrient 

rich. In contrast, the shallow soils at the upper reaches <strong>of</strong> the fan, rich in coarse material 

provide quite unfavourable growth conditions. This is due to their strongly reduced 

water holding capacity <strong>and</strong> thus strong dessication in summer (Prilipko 1954). 


7

-Materials <strong>and</strong> Methods- 

3 Materials <strong>and</strong> Methods 

The field work was done from the end <strong>of</strong> August to the beginning <strong>of</strong> November in 2007. 

On 71 plots (Fig. 3.1) vegetation in different <strong>forest</strong> types, which were determined 

visually before, was sampled. We recorded vegetation, st<strong>and</strong> structure, <strong>and</strong> rejuvenation 

in 59 <strong>of</strong> those sampling plots. In 11 plots we only recorded vegetation <strong>and</strong> rejuvenation. 

Preferably the least influenced plots were chosen, regarding <strong>forest</strong>ry <strong>and</strong> grazing. 

A u f n a h m e f l ä c h e n 

N 

4 k m 

Fig. 3.1 Location <strong>of</strong> the 71 vegetation plots (Map source: (Google Earth 2008)) 

3.1 Vegetation sampling 

Vegetation was recorded in homogeneous 100 m² plots (Fig. 3.2). This design was not 

applicable in four cases, however vegetation was sampled in smaller sized plots (about 

60 m²) adapted to the respective site conditions. The cover <strong>of</strong> each vascular plant 

species occurring in a plot was estimated as well as the cover <strong>of</strong> the herb layer, shrub 

layer, lower tree layer, upper tree layer <strong>and</strong> bare ground. As cover scale we used the ten 

scaled North Carolina vegetation index (Peet et al. 1998). Mean height <strong>of</strong> the tree layer 

was roughly estimated. The vine layer was estimated in a scale ranging from 0 to 3 with 

“0” (no vines), “1” (low presence), “2” (at least every third tree covered) <strong>and</strong> “3”(every 


8

-Materials <strong>and</strong> Methodstree 

covered). Determination <strong>of</strong> the herbarised plant species follows Karjagin (1950). 

Nomenclature follows the Russian reference list (Czerepanov 1995). 

Fig. 3.2 Vegetation sampling design with soil pr<strong>of</strong>ile in the centre 

3.2 Rejuvenation sampling 

Rejuvenation was recorded in five regularly arranged sub-plots <strong>of</strong> 2 x 2 metres within 

the main vegetation plot (Fig. 3.2) resulting in a total sampled area <strong>of</strong> 20 m² per plot. 

Within each plot all juvenile wooden plants in six different classes were counted: 1) 

seedling 2)

-Materials <strong>and</strong> Methodssame 

distance to it. Within the quadrants the following parameters were taken: 1) 

distance to the next tree 2) species <strong>of</strong> that tree 3) its circumference at breast height. 

Every wooden plant with a circumference <strong>of</strong> at least 12 cm was considered as a tree. 

Distance <strong>and</strong> circumference were measured by tape. 

Fig. 3.3 Point Centred Quarter Method (PCQ) - sampling design 

3.4 Site Parameters 

For every sampling plot the coordinates <strong>and</strong> the altitude above sea level were taken with 

the help <strong>of</strong> a Garmin GPS device. Aspect was determined by compass, while inclination 

was estimated visually. A soil sample (0-20cm) was taken in the centre <strong>of</strong> each 100 m² 

plot. Litter height, coarse soil fraction (> 2 mm), fine soil fraction (< 2mm), soil 

moisture <strong>and</strong> carbonate were estimated in the field according to the KA4 (AG Boden 

1996). Grazing intensity was usually estimated using a scale ranging from “0” (no 

grazing) to “3” (intensive grazing) based on the destruction level <strong>of</strong> the vegetation. 

3.5 Data analysis 

Data were stored in the database TurboVEG (version 2.7.4.) <strong>and</strong> statistical analyses 

were run using the program R (version 2.7.2, www.r-project.org). A hierarchical 


10

-Materials <strong>and</strong> Methodsagglomerative 

cluster analysis was applied to the transformed abundance data to 

classify the vegetation. For distance measurement the square root <strong>of</strong> the Bray-Curtis 

coefficient, that has metric properties, was used. 

Table 3.1 Transformed abundance data <strong>of</strong> the North Carolina Index (NCI) 

NCI 0 1 2 3 4 5 6 7 8 9 10 

transformed values 0 0.1 0.5 1.5 3.5 7.5 17.5 37.5 62.5 85 97.5 

As linking procedure the ward’s minimum variance method (Ward 1963) was chosen to 

minimize the squared error for each clustering step. Species occurring in less than three 

plots were not included in this procedure. The socio-ecological species groups were 

arranged manually <strong>and</strong> the plant <strong>communities</strong> were arranged roughly along a moisture 

gradient. The vegetation <strong>communities</strong> were named after two highly frequent species 

present within the community. 

To reveal the correlations with the site parameters, a Detrended Correspondence 

Analysis (DCA) was run with the program PCOrd (version 5.10.) without down 

weighting rare species <strong>and</strong> rescaling axes with 26 segments. All species, occurring less 

than three times in the whole dataset as well as species, that could only be determined to 

genus level were dropped. In order to test if the <strong>forest</strong> types could be distinguished only 

by means <strong>of</strong> their structure, this was repeatedly done using the importance values <strong>of</strong> the 

tree species <strong>and</strong> the same site parameters. 

To reveal successional trends within the <strong>communities</strong> a non-metric multidimensional 

scaling (NMDS) was made. For this purpose from each <strong>of</strong> the 59 sampling plots with 

complete information about canopy structure (PCQ) <strong>and</strong> rejuvenation we created two 

corresponding “sub-plots”. To avoid distortions in the ordination scatter we restricted 

the analysis to 19 tree species that potentially can grow into the tree layers (Table 3.2). 

One sub-plot contained the importance values <strong>of</strong> the PCQ data <strong>and</strong> the second sub-plot 

contained the importance values <strong>of</strong> the respective rejuvenation. To generate importance 

values for the rejuvenation, the numbers <strong>of</strong> individuals within each <strong>of</strong> the six different 

rejuvenation classes were multiplied with a factor (seedling x 0.05 /

-Materials <strong>and</strong> Methods- 

Table 3.2 The following species were included in the NMDS analysis 

1. Acer campestre 

2. Acer velutinum 

3. Alnus barbata 

4. Carpinus caucasica 

5. Carpinus orientalis 

6. Cornus sanguineus 

7. Corylus avellana 

8. Cotinus coggygria 

9. Crataegus spec 

10. Fraxinus exelsior 

11. Mespilus germanica 

12. Morus spec 

13. Populus alba 

14. Populus nigra 

15. Prunus cerasus 

16. Pterocarya pterocarpa 

17. Ulmus foliacea 

18. Quercus iberica 

19. Quercus robur 

To finally allow a common treatment the importance values <strong>of</strong> both the canopy <strong>and</strong> the 

rejuvenation had to be st<strong>and</strong>ardized. St<strong>and</strong>ardization <strong>of</strong> the importance values <strong>and</strong> the 

rejuvenation values was done by dividing each single species value by the highest value 

occurring in the plot. 

The distribution map was created on the basis <strong>of</strong> a satellite image (Google Earth 2008) . 

Indication <strong>of</strong> the vegetation types bases on both the data <strong>of</strong> the sampled vegetation plots 

<strong>and</strong> notifications made during extensive exploration <strong>of</strong> the fan. 

The three riverbed cross-sections were made in different altitudes <strong>of</strong> the fan. For each <strong>of</strong> 

them a GPS value was taken. The l<strong>and</strong>scape is illustrated in a generalised way. 

Proportions do not correspond with the reality. 

The three fan sections are also a generalised representation. They base on both the 

analysed vegetation <strong>communities</strong> with their respective site parameters <strong>and</strong> extensive 

exploration <strong>of</strong> the fan. 


12

-Results- 

4 Results 

4.1 Vegetation <strong>communities</strong> 

71 vegetation plots scattered over the tree-covered area <strong>of</strong> the fan surface were sampled 

(Fig. 3.1). A total <strong>of</strong> 172 species was recorded, among them 30 woody plants (trees <strong>and</strong> 

shrubs) <strong>and</strong> 142 herbs. By means <strong>of</strong> the cluster analysis nine plant <strong>communities</strong> were 

distinguished. Since two <strong>of</strong> them were very similar, they were finally aggregated (Fig. 

4.1). The sampled <strong>communities</strong> comprise pioneer poplar st<strong>and</strong>s close to the river bed, 

two types <strong>of</strong> oak-hornbeam <strong>forest</strong>s with relatively open structure, densely vegetated 

st<strong>and</strong>s <strong>of</strong> ash-wing nut, maple <strong>forest</strong>s with partially high amounts <strong>of</strong> hardly penetrable 

woody vines, white poplar <strong>forest</strong>s along the riverside <strong>and</strong> almost monospecific alder 

<strong>forest</strong>. Mean tree layer heights range from 9 to 28 metres though older individuals <strong>of</strong> 

white poplar grow up to about 50 metres. The mean species number per plot ranges 

from 13 within the open <strong>and</strong> drier <strong>communities</strong> to 24 within the densely vegetated <strong>and</strong> 

moist <strong>communities</strong>. 

6 1 8 

7 

3 5 4 

2 

Fig. 4.1 Dendrogram <strong>of</strong> the plant <strong>communities</strong> based on 71 relevés. Equisetum telmateia-Alnus 

barbata com. (1), Fraxinus excelsior-Pterocarya pterocarpa com. (2), Swida sanguinea-Populus alba 

com. (3), Oplismenus undulatifolius-Acer velutinum com. (4), Quercus robur-Carpinus betulus com. (5), 

Quercus iberica-Carpinus orientalis com. (6), Scleropoa rigida-Populus nigra com. (7), Echinochloa 

crusgalli-Populus alba com. (8) 


13

-Results- 

Table 4.1 Percentage synoptic table <strong>of</strong> the plant <strong>communities</strong>. Further indifferent or rare species are 

shown in Appendix II. Equisetum telmateia-Alnus barbata community. (1), Fraxinus excelsior- 

Pterocarya pterocarpa com. (2), Swida sanguinea-Populus alba com. (3), Oplismenus undulatifolius- 

Acer velutinum com. (4), Quercus robur-Carpinus betulus com. (5), Quercus iberica-Carpinus orientalis 

com. (6), Scleropoa rigida-Populus nigra com. (7), Echinochloa crusgalli-Populus alba com. (8) 

Community 1 2 3 4 5 6 7 8 

Releveś 11 11 10 4 5 16 10 5 

Alnus barbata 100 36 . . . . . 80 

Menta aquatica 45 . . . . . . 20 

Berula erecta 27 . . . . . . . 

Sigesbeckia orientalis 27 9 11 . . . . 20 

Carex elata 27 . . . . . . . 

Lycopus europaeus 64 36 . . . . . 60 

Carex remota 64 45 11 25 20 . . . 

Equisetum telmateia 82 73 33 . . . . 20 

Circea lutetiana 36 27 11 50 . . . . 

Viburnum opulus 36 45 11 . . . . . 

Lonicera caprifolium 73 27 11 25 20 25 . . 

Oplismenus undulatifolius 82 91 44 100 60 . . 20 

Swida sanguinea 91 100 100 100 60 31 10 . 

Euonymus europaeus 45 73 56 100 40 . . . 

Smilax excelsa 91 91 67 75 100 25 . 60 

Hedera helix 82 100 100 100 100 31 10 40 

Geum urbanum 18 36 56 100 40 . . . 

Carpinus betulus 9 73 33 100 80 6 . . 

Morus spec 36 45 78 75 40 6 10 60 

Fraxinus excelsior 27 91 22 75 80 13 . . 

Prunus cerasus 9 64 44 75 60 6 . . 

Acer campestre . 55 11 100 80 19 . . 

Acer velutinum 18 100 33 100 . . . . 

Corylus avellana . 82 33 50 . . . . 

Diospyros lotus 36 64 22 . . . . . 

Pterocarya pterocarpa 18 64 44 25 . 6 . . 

Populus alba 36 18 100 . . 6 50 100 

Ulmus minor 18 18 . 25 60 25 . . 

Quercus robur 9 . 11 . 100 6 10 . 

Urtica dioica . . 22 . 60 . . . 

Phytolacca americana 9 9 33 75 60 . . . 

Mespilus germanica 9 18 33 . 80 75 . . 

Carpinus orientalis 9 . 22 . 20 100 20 . 

Quercus iberica . . . . . 88 20 . 

Cornus mas . . . . . 38 . . 

Sorbus torminalis . . . . 20 31 . . 

Pyracantha coccinea 36 9 44 . . 44 70 . 

Cotinus coggygria . . 22 . . 56 60 20 

Scleropoa rigida 9 . . . . 6 70 20 

Satureja hortensis . . . . . . 40 . 

Kohlrauschia prolifera 9 . . . . . 30 . 

Lactuca spec . . . . . . 40 . 

Daucus carota 9 9 . . . . 50 20 

Phragmitis australis . . . . . . 20 20 

Lotus caucasicus 9 . . . . 6 50 . 

Clematis orientalis . . . . . . 20 20 

Ambrosia artemisifolia . . . . . . 20 20 

Artemisia vulgaris . . . . . . 20 20 

Eleagnus angustifolia . . . . . . 20 20 

Conyza canadensis 18 9 . . . 25 100 80 

Setaria viridis 18 9 . . . 31 90 100 

Bothriochloa ischaemum 9 . . . . 19 70 40 

Populus nigra 9 . 22 . . 6 100 60 

Euphorbia humifusa 9 . . . . . 60 80 

Melilotus <strong>of</strong>ficinale . . . . . 6 30 40 

Echinocloa crusgalli . . . . . . . 80 

Digitaria sanguinea . . . . . . 10 60 

Epilobium hirsutum . . . . . . . 60 

Eragrostis minor 9 . . . . . . 60 

Fimbirstylis sieberiana . . . . . . . 60 

Plantago major . 9 . . . . . 60 

Salix purpurea 9 . . . . . 10 60 

Sonchus oleraceus 9 . . . . . 10 60 

Artemisia annua . . . . . . 10 40 

Digitaria ischaemum 9 . . . . . . 40 

Juncus articulatus . . . . . . . 40 

Persicaria maculata . . . . . . . 40 

Tamarix spec . . . . . . . 40 

Typha angustifolia . . . . . . . 40 


14

-Results- 

In the following chapter species composition <strong>and</strong> st<strong>and</strong> structure will be described in 

greater detail. Table 4.2 gives a short summary <strong>of</strong> the sampling results. 

Table 4.2 Site <strong>and</strong> structure parameters for all <strong>communities</strong> 

Community 1 2 3 4 5 6 7 8 

Releveś 11 11 10 4 5 16 10 5 

Altitude [m] 293 265 256 243 252 404 358 330 

Carbonate [%] (0-20cm) 4 4 5 1 0 1 3 4 

Humidity <strong>of</strong> soil (0-20cm) (Median) 4 2 1 1 2 1 1 3 

Coarse soil [%] (0-20cm) 8 3 13 3 3 31 68 88 

Clay [%] (0-20cm) 29 42 23 52 42 8 3 5 

Silt [%] (0-20cm) 38 40 36 35 49 32 17 14 

S<strong>and</strong> [%] (0-20cm) 33 18 41 13 9 60 81 81 

Litter [cm] 1,3 1,9 2,2 2,1 2,5 2,3 0,2 0,2 

Grazing intensity (Median) 0 0 1 0 3 2 1 1 

Cover herbs [%] 49 59 21 51 2 9 14 19 

Cover shrubs [%] 38 33 45 22 11 46 22 0 

Cover lower tree layer [%] 29 32 33 40 27 50 3 17 

Cover upper tree layer [%] 46 62 60 85 57 2 0 0 

Bare ground [%] 43 35 70 56 88 83 81 83 

Mean height [m] 18 17 24 17 27 10 10 - 

Woody vine density (Median) 2 2 1 2 0 0 0 0 

Structure parameters (PCQ) 

Number <strong>of</strong> trees sampled 220 220 180 80 100 320 60 - 

Number/ha 548 482 792 541 362 979 692 - 

Mean distance [m] 5 5 4 5 5 3 4 - 

BA/ha [m²] 46 37 36 46 61 25 11 - 

Absolute frequency [%] 178 269 231 260 268 215 273 - 

1. Equisetum telmateia-Alnus barbata community 

Fig. 4.2 Importance values <strong>of</strong> Equisetum telmateia-Alnus barbata community (n=11) 

This community is restricted to the middle parts <strong>of</strong> the fan. It is bound to wet <strong>and</strong> loamy 

soils poor in coarse substrates (>2 mm). The mean tree layer height is about 18 metres 

(Fig. 4.7). But some Populus alba individuals grow up to more than 40 metres. The 

average number <strong>of</strong> 23 species per plot is one <strong>of</strong> the highest within the sampled 

<strong>communities</strong> (Fig. 4.7). Vegetation cover is usually high. The mean herb-layer cover 


15

-Results<strong>and</strong> 

also the upper tree layer cover are about 50 %. Woody vines, especially Smilax 

excelsa, Hedera helix <strong>and</strong> Lonicera caprifolium are abundant, creating a dense <strong>and</strong> 

partially hardly penetrable vegetation structure. Alnus barbata is the utterly dominant 

tree species (Fig. 4.2) counting with 428 individuals per hectare, representing nearly 

80 % <strong>of</strong> all present trees. Elder individuals <strong>of</strong> Alnus barbata reach circumferences <strong>of</strong> 

about 250 cm. The total basal area <strong>of</strong> 46 m² per hectare is one <strong>of</strong> the highest <strong>of</strong> all 

<strong>communities</strong> (Fig. 4.6). Typical shrubs are Swida sanguinea <strong>and</strong> Rubus spp., common 

herbs are Equisetum telmateia, Mentha aquatica <strong>and</strong> Carex remota. The grazing 

intensity is low. Main grazers are cattle <strong>and</strong> water buffaloes. Compared to other 

<strong>communities</strong> intensity <strong>of</strong> fuel wood cutting is relatively low. 

Fig. 4.3 Measurements in Equisetum telmateia-Alnus barbata <strong>forest</strong> (left) <strong>and</strong> respective soil pr<strong>of</strong>ile 

(right) 

2. Fraxinus excesior-Pterocarya pterocarpa community 

Fig. 4.4 Importance values <strong>of</strong> Fraxinus excelsior-Pterocarya pterocarpa community (n=11) 


16

-Results- 

This quite dense community stocks on well water supplied sites in the lower parts <strong>of</strong> the 

fan away from stronger disturbances caused by flooding. The coarse soil fraction is 

nearly absent, while clay <strong>and</strong> silt (about 40 % each) clearly dominate the fine-soil 

fraction. The mean height <strong>of</strong> the tree layer is 17 metres. Elder individuals <strong>of</strong> Pterocarya 

pterocarpa <strong>and</strong> Fraxinus excelsior <strong>of</strong>ten grow up to 30 metres <strong>and</strong> higher. Some 

exceptionally large individuals <strong>of</strong> Populus alba can reach circumferences <strong>of</strong> about ten 

metres. With a mean <strong>of</strong> 24 species per plot the species diversity is high. The vegetation 

cover is rather high, particularly the herb layer cover <strong>and</strong> the upper tree layer cover with 

about 60 % each. In contrast bare ground cover is quite low (35 %). Woody vines, 

particularly Hedera helix <strong>and</strong> Smilax excelsa tend to play a major role within this 

community <strong>and</strong> sometimes complicated the tree measurements. This is demonstrated by 

their high frequencies <strong>of</strong> 100 % <strong>and</strong> 90 %, respectively. The absolute tree density 

amounts to 482 individuals per hectare (Fig. 4.6). Carpinus betulus, Fraxinus excelsior 

<strong>and</strong> Pterocarya pterocarpa (Fig 4.4) clearly dominate within the tree layer. Larger 

individuals <strong>of</strong> the former mentioned species count with circumferences <strong>of</strong> 250 cm <strong>and</strong> 

more. The total basal area amounts to 36 m² per hectare (Fig. 4.6), with the above 

mentioned trees also dominating. Further typical wooden plants are Acer velutinum <strong>and</strong> 

Swida sanguinea, common herbs are Equisetum telmateia <strong>and</strong> Oplismenus 

undulatifolius. Grazing intensity is quite low. Logging <strong>and</strong> fuel wood cutting was 

occasionally noted. 

Fig. 4.5 Dense Fraxinus excelsior-Pterocarya pterocarpa <strong>forest</strong> (left) <strong>and</strong> respective soil pr<strong>of</strong>ile (right) 


17

-Results- 

Number <strong>of</strong> trees/ha 

1000 

800 

600 

400 

200 

0 

1 2 3 4 5 6 7 

Community 

70 

60 

50 

40 

30 

20 

10 

0 

1 2 3 4 5 6 7 

Community 

Fig. 4.6 Absolute tree density (left) <strong>and</strong> total basal area (right) per community 

BA/ha [m²] 

Fig. 4.7 Boxplots representing mean number <strong>of</strong> species (top left), mean height <strong>of</strong> tree layer (top right) 

<strong>and</strong> mean s<strong>and</strong> content (soil) per community (bottom) 


18

-Results- 

3. Swida sanguinea-Populus alba community 

Fig. 4.8 Importance values <strong>of</strong> Swida sanguinea-Populus alba community (n=10) 

This community is bound to sites adjacent to the riverbanks in the middle <strong>and</strong> lower 

parts <strong>of</strong> the fan. In the lower parts st<strong>and</strong>s <strong>of</strong> much larger extent exist. Loamy soils are 

favoured, <strong>and</strong> are at least during the hot summer time relatively dry. S<strong>and</strong>y substrates 

obviously dominate (40 %). The coarse soil fraction is relatively low (about 10 %). The 

carbonate fraction here, is the highest in all <strong>communities</strong> (5 %). The mean height <strong>of</strong> the 

tree layer is about 24 metres, which is also one <strong>of</strong> the highest <strong>of</strong> all <strong>communities</strong>. Most 

<strong>of</strong> the Populus alba individuals grow up to 35 metres <strong>and</strong> more, some <strong>of</strong> them reaching 

circumferences <strong>of</strong> about 300 cm. Other species like Ulmus minor <strong>and</strong> Morus spp. are 

much lower in height. The species number is medium-sized with a mean <strong>of</strong> 17 species 

per plot. The vegetation cover is high, especially the shrub layer cover (45 %) <strong>and</strong> the 

upper tree layer cover (60 %). The herb layer cover is relatively low (20 %), 

corresponding with the bare ground cover, which is rather high (70 %). Woody vines, 

especially Hedera helix are common, but not very frequent. The absolute tree density 

amounts to a high value <strong>of</strong> 792 individuals per hectare, particularly caused by Populus 

alba. The total basal area amounts to 36 m² per hectare (Fig. 4.6). Populus alba is the 

most dominant tree species. Further ones are Morus spp. <strong>and</strong> Pterocarya pterocarpa but 

they are much less frequent. Swida sanguinea <strong>and</strong> Rubus spp. are additional typical 

shrub species <strong>of</strong> this community. Geum urbanum is a common herb species. Grazing by 

cattle is common with low to medium intensities. Logging <strong>and</strong> fuel wood cutting 

sometimes occurs. 


19

-Results- 

Fig. 4.9 Populus alba st<strong>and</strong> at the riverbed (left) <strong>and</strong> respective soil pr<strong>of</strong>ile (right) 

4. Oplismenus undulatifolius-Acer velutinum community 

Fig. 4.10 Importance values <strong>of</strong> Oplismenus undulatifolius-Acer velutinum community (n=4) 

This community is bound to the more distant <strong>and</strong> lower parts <strong>of</strong> the fan. It prefers loamy 

to clayey soils. At least during summer time soil humidity is relatively low <strong>and</strong> soil 

structure is very solid, which makes it rather difficult to dig. Dominant fractions within 

the fine soil are clay <strong>and</strong> silt with mean values <strong>of</strong> 50 % <strong>and</strong> 35 %, respectively. The 

coarse soil fraction is nearly absent. The carbonate content is very low (1 %). The mean 

tree layer height is about 17 metres, although older Acer velutinum <strong>and</strong> Carpinus 

betulus trees grow up to more than 30 metres. The mean species number per plot is 20, 

which is a medium value among the sampled <strong>communities</strong>. The vegetation cover is 

quite high, especially the upper tree layer cover (85 %) <strong>and</strong> the herb layer cover (50 %). 


20

-Results- 

Woody vines, such as Hedera helix <strong>and</strong> Smilax excelsa are very frequent within the 

community. The value for the absolute tree density is with 541 individuals per hectare 

not conspicuous, but in turn the total basal area amounts to 46 m² per hectare, which is 

the second largest value among the <strong>communities</strong> (Fig. 4.6.) Acer velutinum is the most 

important tree species, reaching maximal circumferences <strong>of</strong> about 270 cm. Further 

common tree species are Carpinus betulus <strong>and</strong> Fraxinus excelsior. Although not <strong>of</strong> 

greater importance, Acer campestre is a very typical species within this community. 

Further typical species are Euonymus europaeus <strong>and</strong> Prunus cerasus within the shrub 

layer, Oplismenus undulatifolius <strong>and</strong> Geum urbanum within the herb layer. The grazing 

intensity is relatively low. Some marked trees, scattered tree stubs <strong>and</strong> the occasional 

elimination <strong>of</strong> strong Hedera helix individuals, climbing up valuable tree trunks, 

demonstrate the human impact. 

Fig. 4.11 St<strong>and</strong> <strong>of</strong> Acer velutinum <strong>and</strong> Fraxinus excelsior (left) <strong>and</strong> respective soil pr<strong>of</strong>ile (right) 


21

-Results- 

5. Quercus robur-Carpinus betulus community 

Fig. 4.12 Importance values <strong>of</strong> Quercus robur-Carpinus betulus community (n=5) 

This community is located in the upper <strong>and</strong> middle parts <strong>of</strong> the fan, where it presents a 

remarkable open structure with predominantly old, mighty individuals <strong>of</strong> Quercus 

robur, smaller ones <strong>of</strong> Carpinus betulus <strong>and</strong> a nearly absent under storey. It is bound to 

slightly moist soils. Carbonate is completely absent. Clayey (40 %) <strong>and</strong> silty (50 %) 

fractions clearly dominate within the substrate, the portion <strong>of</strong> coarse material is very 

low. There are few differences in tree heights. The mean tree layer height amounts to 27 

metres, the highest value <strong>of</strong> all sampled <strong>communities</strong>. Species diversity is medium, with 

a mean number <strong>of</strong> 18 species per plot. The vegetation cover is not very high, bare soil is 

dominating the ground. Woody vines are nearly absent. The absolute tree density with 

362 individuals per hectare is compared with the other <strong>communities</strong> strikingly low 

(Fig.4.6). In turn it is the community with the highest total basal area (61 m² per 

hectare). Quercus robur <strong>and</strong> Carpinus betulus are the most prominent tree species. 

Particularly Quercus individuals are huge-grown <strong>and</strong> their circumferences are <strong>of</strong>ten 

more than 300 cm. Further common tree species are Fraxinus excelsior, Acer campestre 

<strong>and</strong> Mespilus germanica. The abundance <strong>of</strong> disturbance indicator species like Urtica 

dioica <strong>and</strong> Phytolacca americana is conspicuous. The community is highly influenced 

by men <strong>and</strong> livestock indicated by the common tree stubs <strong>and</strong> the almost completely 

absence <strong>of</strong> under storey <strong>and</strong> rejuvenation except under thorny vegetation <strong>of</strong> Smilax 

excelsa. 


22

-Results- 

Fig. 4.13 Quercus robur-Carpinus betulus community (left) with a nearly absent under storey <strong>and</strong> 

respective soil pr<strong>of</strong>ile (right) 

6. Quercus iberica-Carpinus orientalis community 

Fig. 4.14 Importance values <strong>of</strong> Quercus iberica-Carpinus orientalis community (n=16) 

With its pronounced tolerance for drier soil conditions, this low-grown community is 

restricted to the upper reaches <strong>of</strong> the fan. The coarse soil fraction is rather high (30 %). 

Within the fine soil s<strong>and</strong>y fractions clearly predominate (60 %), while carbonate is 

nearly absent (1 %). The mean tree layer height is about 10 metres. Even the highest 

Quercus iberica do not grow up to more than 20 metres. Shrubs <strong>and</strong> low grown trees 

dominate the scenery. It is a rather species-poor community with a mean <strong>of</strong> 13 species 

per plot. Shrub layer cover <strong>and</strong> lower tree layer cover are relatively high with 45 <strong>and</strong> 

50 %, respectively. Woody vines are completely absent. In contrast to the above 


23

-Resultsdescribed 

Quercus robur–Carpinus betulus community the absolute tree density is the 

highest among the <strong>communities</strong> with 979 individuals per hectare, whereas the total 

basal area is strikingly low with only 25 m² per hectare (Fig.4.6). Carpinus betulus is 

the most abundant species with 581 individuals per hectare, but the far less frequent 

Quercus iberica is responsible for nearly half <strong>of</strong> the total basal area portion. Further 

important accompanying species are Mespilus germanica, Ligustrum vulgare, Cotinus 

coggygria, Crataegus spp. <strong>and</strong> Rosa spp. within the shrub layer. Herbal vegetation 

dominated by Setaria viridis <strong>and</strong> Conyza canadensis occurs scattered, while bare 

ground fraction amounts to a considerably high value (83 %). The grazing intensity is 

moderate. Principal grazer are sheep. The human impact is obvious <strong>and</strong> particularly 

visible at the <strong>of</strong>ten multi-stemmed Carpinus orientalis <strong>and</strong> stubs <strong>of</strong> recently logged 

Quercus iberica trees. 

Fig. 4.15 Quercus iberica-Carpinus orientalis <strong>forest</strong> with multi stemmed Carpinus orientalis indicating 

regularly fuel wood cut (left) <strong>and</strong> soil pr<strong>of</strong>ile (right) 


24

-Results- 

7. Scleropoa rigida-Populus nigra community 

Fig. 4.16 Importance values <strong>of</strong> Scleropoa rigida-Populus nigra community (n=3) 

This community comprises both very open, pioneer st<strong>and</strong>s as well as st<strong>and</strong>s, which are 

to a higher degree vertically structured . It is a rather open community with a large 

number <strong>of</strong> relatively small sized trees. Restricted to the riverbanks <strong>and</strong> open gravel beds 

<strong>of</strong> the upper reaches <strong>of</strong> the fan it is subjected to frequent disturbances by flooding. The 

substrate is considerably rough structured with a mean coarse soil fraction <strong>of</strong> 68 % <strong>and</strong> 

a predomination <strong>of</strong> s<strong>and</strong>y fractions (80 %) within the fine soil. Due to the high degree <strong>of</strong> 

disturbance the plants have to deal with strongly differing soil moisture conditions 

during the course <strong>of</strong> the year. At least in summer soil moisture is relatively low. The 

mean tree layer hight is about ten metres (pioneer st<strong>and</strong>s excluded). Some elder Populus 

nigra or Populus alba individuals grow up to about 20 metres, both reaching maximal 

circumference values <strong>of</strong> about 300 cm. With a mean <strong>of</strong> 18 species per plot it is a 

moderately species rich community. Vegetation cover is usually very low, <strong>and</strong> bare 

ground predominates (80 %). The absolute tree density amounts to a considerably high 

value <strong>of</strong> 692 individuals per hectare (Fig. 4.6), though the amount <strong>of</strong> the total basal area 

is strikingly low (11 m² per hectare). Particularly Populus nigra <strong>and</strong> Cotinus coggygria 

turn out to be the most important woody species. Cotinus coggygria occurs with a large 

number <strong>of</strong> individuals <strong>and</strong> Populus nigra with less but larger sized individuals. 

Pyracantha coccinea, Conyza canadensis, Setaria viridis <strong>and</strong> Scleropoa rigida are 


25

-Resultsfurther 

typical species <strong>of</strong> this community. The grazing intensity is relatively low. Main 

grazers are sheep. 

Fig. 4.17 Populus nigra st<strong>and</strong> with Cotinus coggygria in the front (left), rejuvenation <strong>of</strong> Populus nigra 

(right) <strong>and</strong> respective soil pr<strong>of</strong>ile (centre) - all close to the riverbed 

8. Echinocloa crus-galli-Populus alba community 

This pioneer-community is located in the middle <strong>and</strong> lower parts <strong>of</strong> the fan. It is bound 

to moist <strong>and</strong> open, not too sunny river beds with a high amount <strong>of</strong> coarse material. 

Thus, the mean coarse soil fraction is about 90 %. Soil-carbonate is relatively high 

(4 %). Within the fine-soil s<strong>and</strong> is clearly dominating (80 %). The mean <strong>of</strong> 24 species 

per plot is one <strong>of</strong> the highest values <strong>of</strong> all surveyed <strong>communities</strong>. Herbs cover about 

20 % <strong>of</strong> the area, but the major part is not covered by vegetation (85 %). Higher trees 

are completely absent, only sometimes trees <strong>of</strong> adjacent <strong>communities</strong> partly contributed 

to the tree layer cover. Dominant woody plants apart from Populus alba are young 

individuals <strong>of</strong> Alnus barbata <strong>and</strong> Salix purpurea, whereas Echinochloa crusgalli, 

Eragrostis minor, Euphorbia humifusa, Fimbristylis sieberiana <strong>and</strong> a number <strong>of</strong> other 

annual plants dominate within the herb layer. Grazing is common, but with a low 

intensity, mainly by sheep <strong>and</strong> cattle. 


26

-Results- 

Fig. 4.18 Rejuvenation sampling <strong>of</strong> Populus alba in the riverbed (left) <strong>and</strong> respective soil pr<strong>of</strong>ile (right) 

4.2 Rejuvenation 

Within this chapter the rejuvenation <strong>of</strong> ten tree species will be described. The following 

species turned out to be the most important ones. They are treated in alphabetical order: 

Acer velutinum 

This species displays considerable rejuvenation rates at the middle <strong>and</strong> lower parts <strong>of</strong> 

the fan. It apparently favours humid soil conditions, a nearly absent coarse soil fraction 

<strong>and</strong> loamy to clayey substrate conditions. It occurs under relatively high canopy cover 

on rather well vegetated ground. These conditions are given within the Equisetum 

telmateia-Alnus barbata community, the Fraxinus excelsior-Ptereocarya pterocarpa 

<strong>and</strong> particularly within the Oplismenus undulatifolius-Acer velutinum community (Fig. 

4.19 <strong>and</strong> Fig. 4.20). The almost completely absence <strong>of</strong> juvenile plants above 50 cm is 

remarkably. Hence, only a low amount <strong>of</strong> juveniles grow out <strong>of</strong> the reach <strong>of</strong> livestock (> 

150 cm). 

Alnus barbata 

This species shows higher rejuvenation rates only on the middle <strong>and</strong> lower parts <strong>of</strong> the 

fan within the Equisetum telmateia-Alnus barbata community <strong>and</strong> the pioneer 

community <strong>of</strong> Populus alba (Fig. 4.19 <strong>and</strong> Fig. 4.21). Rather open, light sites, wet soil 

conditions <strong>and</strong> a relatively high carbonate content are preferred. However, substrate 


27

-Resultsproperties 

differ considerably, ranging from very coarse <strong>and</strong> gravelly to pronounced 

loamy conditions. Almost all sampled juvenile plants were less than 50 cm in height <strong>and</strong> 

thus within the range <strong>of</strong> livestock. 

Carpinus betulus <strong>and</strong> Carpinus orientalis 

Since these two species are hardly distinguishable from each other within their juvenile 

stages they will be treated together. Only within the Quercus iberica-Carpinus 

orientalis community a considerable amount <strong>of</strong> “older” juveniles could be certainly 

determinated as Carpinus orientalis, growing on relatively dry, open <strong>and</strong> rough textured 

soils. Apart from this, Carpinus spp. was present in all <strong>communities</strong> which provided 

fine textured <strong>and</strong> well water supplied soils. Open <strong>and</strong> frequently disturbed st<strong>and</strong>s are 

avoided. The recorded rejuvenation was seldom higher than 150 cm. 

Fraxinus excelsior 

This species mainly rejuvenates on the middle <strong>and</strong> lower parts <strong>of</strong> the fan. Loamy to 

clayey, principally humid soils with a low coarse soil fraction are favoured. The 

vegetation cover, in particular the tree layer cover is relatively high. Rejuvenation is 

abundant within the Equisetum telmateia-Alnus barbata community, the Oplismenus 

undulatifolius-Acer velutinum community, the Quercus robur-Carpinus betulus 

community <strong>and</strong> within the Fraxinus excelsior-Pterocarya pterocarpa community (Fig. 

4.19 <strong>and</strong> Fig. 4.20). Mostly, rejuvenation does not grow above 100 cm. 

Populus alba 

Rejuvenation predominantly occurs on the middle <strong>and</strong> lower parts <strong>of</strong> the fan but also in 

its upper parts. It is confined to the vicinity <strong>of</strong> the river bed or small channels. 

Constantly humid soil conditions <strong>and</strong> open, not too sunny places are favoured. Substrate 

properties may range from very coarse to loamy underground. Due to its propagation by 

means <strong>of</strong> root suckers growing from the lateral roots sometimes as far as 20-30 metres 

from the trunk, identification <strong>of</strong> individuals was sometimes difficult. Under drier soil 

conditions plants display much less vitality. It was frequently recorded within the 

Equisetum telmateia-Alnus barbata <strong>and</strong> the Echinochloa crusgalli-Populus alba 


28

-Resultscommunity 

(Fig. 4.19 <strong>and</strong> Fig. 4.21). Individuals larger than 150 cm were scarcely 

found. 

Populus nigra 

Black poplar rejuvenation frequently occurs in, or adjacent to the river bed on the upper 

<strong>and</strong> mid-fan. Juvenile plants were numerously found within the riverbed area, but were 

already lacking in older Populus nigra st<strong>and</strong>s, which are to a higher extent vertically 

structured (Fig. 4.21). Open, sunny places with coarse <strong>and</strong> gravely soil substrate 

properties <strong>and</strong> thus relatively dry soil conditions are preferred. Moister soils are less 

densely populated. Vegetation cover is usually very low. What was said about the root 

suckering habit <strong>of</strong> Populus alba, also applies to Populus nigra. A considerably high 

amount <strong>of</strong> juvenile plants were higher than 150 cm <strong>and</strong> thus grown out <strong>of</strong> the reach <strong>of</strong> 

livestock. 

Pterocarya pterocarpa 

Loamy, clayey <strong>and</strong> principally moist substrates on lower fan sites are preferred, though 

rejuvenation on relatively dry soils was also noticed. It was frequently found on sites 

with rather high vegetation cover, particularly herb <strong>and</strong> upper tree layer cover. Since it 

is also a clonal propagating species, individual counting was sometimes complicated. 

Rejuvenation occurs mainly within the Fraxinus excelsior-Pterocarya pterocarpa <strong>and</strong> 

partly within the Oplismenus undulatifolius-Acer velutinum community (Fig. 4.19 <strong>and</strong> 

Fig. 4.20). 

Quercus iberica <strong>and</strong> Quercus robur 

The two oak species have to be treated together as Quercus spp. because <strong>of</strong> the 

difficulties in identification <strong>of</strong> juvenile Quercus. Rejuvenation <strong>of</strong> Quercus spp. was 

basically found at the upper parts <strong>of</strong> the fan within the Quercus iberica-Carpinus 

orientalis community (Fig. 4.20). Sunny sites, sparsely vegetated ground <strong>and</strong> relatively 

coarse, s<strong>and</strong>y substrates are favoured. Within the other <strong>communities</strong> it was sparsely 

found. All plants were less than 50 cm high <strong>and</strong> thus within the reach <strong>of</strong> livestock. 


29

-Resultsa) 

cm 

b) 

cm 

c) 

cm 

Fig. 4.19 Rejuvenation classes: 1 seedling 2

-Resultsd) 

cm 

e) 

cm 

f) 

cm 


-Resultsg) 

cm 

h) 

cm 

cm 


-Results- 

4.3 Ordination results 

5 

7 

6 

8 

1 

2 

4 

3 

5 

2 

6 

7 

1 

4 

8 

3 

Fig. 4.22 Ordination diagram (DCA) <strong>of</strong> 71 sample plots generated on the basis <strong>of</strong> presence <strong>and</strong> 

abundance <strong>of</strong> 85 plant species. Eight plant <strong>communities</strong> were discriminated. Vectors indicate gradients in 

site parameters. Equisetum telmateia-Alnus barbata com. (1), Fraxinus excelsior-Pterocarya pterocarpa 

com. (2), Swida sanguinea-Populus alba com. (3), Oplismenus undulatifolius-Acer velutinum com. (4), 

Quercus robur-Carpinus betulus com. (5), Quercus iberica-Carpinus orientalis com. (6), Scleropoa 

rigida-Populus nigra com. (7), Echinochloa crusgalli-Populus alba com. (8) 

Table 4.3 Correlation between selected environmental parameters <strong>and</strong> DCA axes 

total inertia 7,92 

Axis 1 Axis 2 Axis 3 

length <strong>of</strong> gradient 4,27 3,36 2,44 

eigenvalue 0,92 0,60 0,35 

correlation coefficient r r r 

altitude 0,695 0,137 0,233 

moisture -0,471 -0,082 0,192 

carbonate -0,382 -0,357 -0,007 

skeleton 0,463 -0,157 0,480 

clay -0,606 0,179 -0,322 

silt -0,241 0,201 -0,324 

s<strong>and</strong> 0,559 -0,242 0,413 


33

-Results- 

The DCA analysis (Fig. 4.22) revealed a clear differentiation <strong>of</strong> the eight plant 

<strong>communities</strong> along the first two ordination axes. Correlation is strong between the first 

axis <strong>and</strong> the site parameters altitude, soil structure <strong>and</strong> soil moisture (Table 4.3). The 

second axis shows no significant correlations with one <strong>of</strong> the measured site parameters. 

3 

7 

1 

6 

2 

4 

5 

Fig. 4.23 Ordination diagram (DCA) <strong>of</strong> 59 sample plots generated on the basis <strong>of</strong> different canopy 

importance values. Seven <strong>forest</strong> <strong>communities</strong> are discriminated. Vectors indicate gradients in site 

parameters. Equisetum telmateia-Alnus barbata com. (1), Fraxinus excelsior-Pterocarya pterocarpa com. 

(2), Swida sanguinea-Populus alba com. (3), Oplismenus undulatifolius-Acer velutinum com. (4), 

Quercus robur-Carpinus betulus com. (5), Quercus iberica-Carpinus orientalis com. (6), Scleropoa 

rigida-Populus nigra com. (7) 

Table 4.4 Correlation between selected environmental parameters <strong>and</strong> DCA axes 

total inertia 5,0 

Axis 1 Axis 2 Axis 3 

length <strong>of</strong> gradient 4,22 3,98 3,42 

eigenvalue 0,86 0,55 0,31 

correlation coefficient r r r 

altitude 0,646 0,044 -0,150 

moisture -0,536 0,052 -0,073 

carbonate -0,385 0,369 -0,091 

skeleton 0,543 0,255 0,114 

clay -0,510 -0,266 0,218 

silt -0,249 -0,305 0,009 

s<strong>and</strong> 0,541 0,392 -0,171 


34

-Results- 

The second ordination bases on importance values <strong>of</strong> 59 relevés <strong>and</strong> shows a similar 

pattern as DCA I (Fig. 4.22). All <strong>communities</strong> appear clearly differentiated along the 

first two ordination axes. Altitude, soil moisture <strong>and</strong> soil structure are strongly 

correlated to the first axis (Table 4.4), whereas the second axis shows no significant 

correlation to any <strong>of</strong> the measured site parameters. 

4.4 Ordination results (Successional trends) 

Rejuvenation has already been mentioned in terms <strong>of</strong> single species recruitment (see 

chapter 4.2). To additionally reveal successional trends within the <strong>communities</strong> a 

NMDS analysis was made. 

5 

4 

6 

2 

1 

7 

3 

Fig. 4.24 Ordination diagram (NMDS) comparing tree species composition <strong>and</strong> respective <strong>regeneration</strong> 

within 59 sampling plots. Same symbols represent same <strong>communities</strong>. Canopy importance is indicated in 

black, rejuvenation importance in red. Equisetum telmateia-Alnus barbata com. (1), Fraxinus excelsior- 

Pterocarya pterocarpa com. (2), Swida sanguinea-Populus alba com. (3), Oplismenus undulatifolius- 

Acer velutinum com. (4), Quercus robur-Carpinus betulus com. (5), Quercus iberica-Carpinus orientalis 

com. (6), Scleropoa rigida-Populus nigra com. (7) 


35

-Results- 

The NMDS-analysis revealed a good differentiation <strong>of</strong> the seven canopy plots similar to 

that <strong>of</strong> the second DCA (Fig. 4.23), with only few outliers. They are arranged both 

along the first <strong>and</strong> along the second axis <strong>of</strong> the diagram. Rejuvenation reveals a trend 

towards the upper <strong>and</strong> particularly the upper right part <strong>of</strong> the diagram. 


36

-Discussion- 

5 Discussion 

5.1 Vegetation zonation <strong>and</strong> site conditions 

The observed alluvial fan is a rather diverse structured l<strong>and</strong>form generated by erosion 

<strong>and</strong> deposition due to recurring inundation. The map <strong>of</strong> the vegetation <strong>communities</strong> 

(Fig. 5.1) shows a characteristic pattern <strong>of</strong> <strong>forest</strong> <strong>communities</strong> apparently following an 

altitudinal gradient. 

5 0 0 m 

Q 

a k h 

1 

b 

2 

a 

4 0 0 m 

3 

3 0 0 m 

N 

4 k m 

2 0 0 m 

r i v e r b e d 

Q u e r c u s i b e r i c a - C a r p i n u s o r i e n t a l i s 

Q u e r c u s r o b u r - C a r p i n u s b e t u l u s 

F r a x i n u s e x c e l s i o r - P t e r o c a r y a p t e r o c a r p a 

S w i d a s a n g u i n e a - P o p u l u s a l b a 

E q u i s e t u m t e l m a t e i a - A l n u s b a r b a t a 

O p l i s m e n u s u n d u l a t i f o l i u s - A c e r v e l u t i n u m 

r o a d 

S c l e r o p o a r i g i d a - P o p u l u s n i g r a 

r a i l r o a d 

P u n i c a g r a n a t a - P a l i u r u s s p i n a - c h r i s t i c r o s s - s e c t i o n 

v e r y l a r g e P o p u l u s a l b a 

c o n c r e t e e m b a n k m e n t 

a g r i c u l t u r e , p a s t u r e o r s m a l l f o r e s t p a t c h e s 

s e t t l e m e n t 

Fig. 5.1 Rough map <strong>of</strong> the alluvial fan <strong>and</strong> the distribution <strong>of</strong> the <strong>forest</strong> <strong>communities</strong> based on a satellite 

image (Google Earth 2008) Mixed st<strong>and</strong>s are depicted striped. The two fan cross sections are indicated by 

bold dashed lines ( ) with a = upper fan, b =lower fan. The three riverbed cross sections are indicated 

with 1 = upper fan, 2 = mid-fan, 3 =lower fan. 

This gradient was considered highly significant within both conducted DCA analyses. 

Further significant gradients revealed by both DCA were soil moisture <strong>and</strong> substrate 

composition. The correlation <strong>of</strong> higher located sites to high fractions <strong>of</strong> coarse material 


37

-Discussion<strong>and</strong> 

lower sites to high fractions <strong>of</strong> fine material due to decreasing discharge power 

towards the more distant parts is characteristic for alluvial fans (Ahnert 1996) <strong>and</strong> has 

also been proven within this study (Fig. 5.2). The figure shows the occurrence <strong>of</strong> 

extremely coarse material (87,5 %) also in mid-altitudes. This is due to the underlying 

sampling plots, which were located within or very close to the river bed, where coarse 

material gets far more downstream than outside. Soil moisture is <strong>of</strong>ten correlated with 

substrate structure. Coarse soils with their large pores allow a relatively rapid water 

infiltration <strong>and</strong> provide a rather low water holding capacity. Hence, particularly in 

summer desiccation <strong>of</strong> these soil types is relatively quick. In turn finer soils allow a 

slower water infiltration but provide a much higher water holding capacity because <strong>of</strong> 

their small pores <strong>and</strong> the resulting stronger soil adhesion (Scheffer & Schachtschabel 

2002). Hence, one should expect the good water supplied soils in the lower <strong>and</strong> more 

distant parts <strong>of</strong> the fan. This has also been proven; though already in the mid-fan 

extremely wet soils occurred (Fig. 5.2). 

Fig. 5.2 Boxplots representing correlations between altidude <strong>and</strong> coarse soil (left) <strong>and</strong> altitude <strong>and</strong> 

humidity (right) 

They were mainly covered by Equisetum telmateia-Alnus barbata <strong>forest</strong>. This is 

certainly due to emerging groundwater. Finally, in line with Blom & Voesenek (1996) 

<strong>and</strong> Ellenberg (1988) hydrological conditions, or more precisely flooding frequency <strong>and</strong> 

duration turn out to be most important for the zonation <strong>of</strong> the vegetation (Kramer et al. 

2008). 


38


N 

E 

S 

W 

6 0 0 m 

4 0 0 m 

S l o p e 

2 0 0 m 

V a l l e y F l o o r 

1 0 k m 

S c l e r o p o a r i g i d a - P o p u s n i g r a 

c o m m u n i t y 

Q u e r c u s i b e r i c a - C a r p i n u s o r i e n t a l i s 


E q u i s e t u m t e l m a t e i a - A l n u s b a r b a t a 


S w i d a s a n g u i n e a - P o p u l u s a l b a 


C o t i n u s c o g g y g r i a s h r u b l a n d Q u e r c u s r o b u r - C a r p i n u s b e t u l u s 


F r a x i n u s e x c e l s i o r - P t e r o c a r y a p t e r o c a r p a 


O p l i s m e n u s u n d u l a t i f o l i u s - A c e r v e l u t i n u m 


w a t e r f l o w r o c k s , s t o n e s g r a v e l , s a n d s i l t , c l a y 

Fig. 5.3 Longitudial section <strong>of</strong> the fan 

Direct effects <strong>of</strong> flooding include reduced oxygen exchange <strong>and</strong> hampered soil water 

movement (Hughes et al. 2001) uprooting (Karrenberg et al. 2003) as well as burial by 

sediments (Marigo et al. 2000). Indirect effects are the creation <strong>of</strong> bare substrates due to 

sedimentation (Johnson 2000), decreased vegetation cover <strong>and</strong> increased nutrient supply 

(Thoms 2003). The response to the above mentioned factors varies among species. The 

s<strong>of</strong>twood species Salix spp. <strong>and</strong> Populus spp. tend to be better adapted to flooding than 

the hardwood broad leafed deciduous species Quercus spp, Ulmus spp. <strong>and</strong> Fraxinus 

spp. (Vreugdenhil et al. 2006). In Europe, this results in a typical zonation <strong>of</strong> the 

s<strong>of</strong>twood species Salix spp. <strong>and</strong> Populus spp. near the main channel <strong>and</strong> the hardwood 

species Quercus spp., Ulmus spp. as well as Fraxinus spp. dominating the less 

frequently flooded sites (Ward et al. 2002; Ellenberg 1988). These observations could 

be affirmed within this study, though the s<strong>of</strong>twood zone was <strong>of</strong>ten narrow <strong>and</strong> 

sometimes completely lacking, possibly due to already reduced river dynamics. Both 

poplar species were obviously restricted to bare soils in, or adjacent to the open riverbed 

where flood dynamic is greatest. Populus nigra prefers the upper sites (Fig. 5.3 , Fig. 

5.4 <strong>and</strong> Fig. 5.6), obviously better adapted to survive drier periods in summer. Glenz et 

al. (2006) indicate its flooding tolerance with high (4) (Table 5.1). 


39


4 2 0 m 

N W S E 

3 7 0 m 

G R A V E L 

6 k m 

Q u e r c u s i b e r i c a - C a r p i n u s o r i n e n t a l i s c o m m u n i t y 

C o t i n u s c o g g y g r i a - s h r u b l a n d 3 7 0 m A l t i t u d e a . s . l . 

S c l e r o p o a r i g i d a - P o p u l u s n i g r a c o m m u n i t y P u n i c a g r a n a t a - P a l i u r u s s p i n a - c h r i s t i 

g r a s s l a n d 

r i v e r b e d ( g r a v e l ) 

Fig. 5.4. Cross section <strong>of</strong> the upper fan 

But due to the high river dynamics which expose it to the above mentioned direct 

flooding effects Populus nigra st<strong>and</strong>s are regularly damaged or completely destroyed by 

floods. In the case <strong>of</strong> channel avulsion st<strong>and</strong>s might be cut <strong>of</strong>f by sufficient water 

supply. Trees then display much less vitality <strong>and</strong> those sites become populated by 

drought-adapted shrubs like Cotinus coggygria <strong>and</strong> Crataegus spp., later by trees like 

Carpinus orientalis <strong>and</strong> Quercus iberica. Flooding tolerance <strong>of</strong> Populus alba is 

somewhat lower (3) (Table 5.1). It displaces Populus nigra at permanently humid sites 

on the mid- <strong>and</strong> lower fan (Fig 5.5 <strong>and</strong> Fig. 5.7). 

Table 5.1 Flooding tolerance classes <strong>of</strong> 65 European tree <strong>and</strong> shrub species based on cluster 

analysis <strong>and</strong> expert knowledge (Glenz et al. 2006) 


40


3 3 0 m 

N W S E 

2 8 0 m 

l o a m y g r o u n d 

G R A V E L 

l o a m y g r o u n d 

Fig. 5.5 Cross section <strong>of</strong> the lower fan 

Soil texture seems to have a lower importance. It may range from coarse to fine. The 

slight correlation between Populus alba st<strong>and</strong>s <strong>and</strong> a relatively high carbonate content, 

revealed by both DCA is rather due to its restriction to young, bare <strong>and</strong> thus more 

carbonate rich soils than to its preference for carbonate (Rothmaler et al. 2002). Neither 

Populus alba nor Populus nigra seem to be good competitors in mixed st<strong>and</strong>s. While 

dominance within their own <strong>communities</strong> is explicit (Fig. 4.8. <strong>and</strong> Fig 4.16), they hardly 

appear within the other <strong>communities</strong>. This is also mentioned by Johnson (2000). While 

younger stages <strong>of</strong> Populus alba <strong>forest</strong> are light <strong>and</strong> fairly monospecific, few old <strong>and</strong> 

huge Populus alba individuals within the Fraxinus excelsior-Pterocarya pterocarpa <strong>and</strong> 

the Oplismenus undulatifolius-Acer velutinum community seem to be remnants <strong>of</strong> the 

aforementioned open riverbed <strong>communities</strong>, still surviving because <strong>of</strong> lacking light 

competition within the upper tree layer. The largest <strong>of</strong> them grow up to 50 metres <strong>and</strong> 

count with circumferences <strong>of</strong> about ten metres. The other <strong>communities</strong> are located with 

more distance to the riverbed, where the direct flood impact is lower <strong>and</strong> sedimentation 

processes predominate due to the decelerated water flow. The Quercus iberica- 

Carpinus orientalis community occupies large parts <strong>of</strong> the upper fan down to about 350 

metres (Fig. 5.1 <strong>and</strong> Fig. 5.3). According to the DCA it is the community with the 

highest correlation to higher located fan-sites, which means with the best adaptations to 

survive drier conditions, like in the fairly hot summer. Different authors describe similar 

<strong>forest</strong>s st<strong>and</strong>s in former Yugoslavia (Dinic et al. 1980) <strong>and</strong> also in Azerbaijan as 

thermophile <strong>forest</strong> st<strong>and</strong>s adapted to relatively dry conditions (Missouri Botanical 

Garden 2009). 


41


5 0 m 

2 m 

Fig. 5.6 Cross section <strong>of</strong> the riverbed (upper fan) 

Of all <strong>communities</strong>, due to their elevated location <strong>and</strong> their coarse substrate structure, 

they are probably the least effected community by long lasting inundations. Further 

downward follows a relatively small belt <strong>of</strong> a second oak-hornbeam community, 

composed <strong>of</strong> Quercus robur <strong>and</strong> Carpinus betulus (Fig. 5.1 <strong>and</strong> Fig. 5.3). The DCA 

revealed a preference for rather fine textured soils <strong>and</strong> slightly moist soil conditions. 

While Quercus robur is known as a typical hardwood species <strong>of</strong> riparian <strong>forest</strong>s, 

capable to endure longer lasting inundations (Ellenberg 1988; Glenz et al. 2006; Kramer 

et al. 2008), Carpinus betulus is not. Its flooding tolerance is indicated with only 2 

(Table 5.1). Its frequent appearance in most <strong>of</strong> the sampled <strong>communities</strong> could be due to 

disrupted water inundations for a longer period <strong>of</strong> time <strong>and</strong> a subsequent drop <strong>of</strong> ground 

water table as observed in the river Rhine basin <strong>and</strong> in the Cahnov-Soutuk <strong>forest</strong> in the 

Czech Republic (Schnitzler 1994; Janik et al. 2008). Permanent springs reveal emerging 

ground water conditions, which favour the establishment <strong>of</strong> Alnus barbata <strong>forest</strong> st<strong>and</strong>s 

in the mid-fan. The abundant presence <strong>of</strong> Equisetum telmateia strongly supports this, 

since it is an indicator for emerging ground water, which is rich in carbonate. Due to its 

outst<strong>and</strong>ing adaptations to survive a long time under anoxic conditions (Glenz et al. 

2006) Alnus barbata dominance within the tree layer is pronounced. 


42


7 0 m 

2 m 

Fig. 5.7 Cross section <strong>of</strong> the river bed (mid-fan) 

Due to the relatively high abundance <strong>of</strong> other species a few plots appear a little more 

distant within the second DCA. They are already representing transition-types to other 

<strong>communities</strong>. The remote <strong>and</strong> less frequently inundated parts <strong>of</strong> the fan are mainly 

covered by <strong>of</strong>ten mixed Fraxinus excelsior-Pterocarya pterocarpa <strong>and</strong> Oplismenus 

undulatifolius-Acer velutinum <strong>communities</strong> (Fig. 5.1 <strong>and</strong> Fig. 5.8). While Fraxinus 

excelsior is recognized as typical hardwood species <strong>of</strong> European riparian <strong>forest</strong>s 

(Ellenberg 1988; Ward et al. 2002) with a flooding tolerance value <strong>of</strong> 3 (Table 5.1), few 

information is available for Pterocarya pterocarpa. Mitchel (1974) describes it as 

typical species <strong>of</strong> riverine <strong>forest</strong>s, able to survive short-term inundations <strong>and</strong> bound to 

humid soil conditions. Stuchlik & Kvavadze (1998) describe it as predominant species 

in the floodplain <strong>forest</strong>s <strong>of</strong> the Alasan river in Georgia; the river that catches the water 

<strong>of</strong> the Kurmukcay river. Since both species have a moderate flooding tolerance, <strong>and</strong> a 

preference for constantly humid soil conditions the restriction to the lower fan parts is 

consequential. Schnitzler (1994) mentions the occupation <strong>of</strong> the ecological niche <strong>of</strong> 

Ulmus minor by Fraxinus excelsior in the Rhine Valley following the quasi 

disappearance <strong>of</strong> the typical hardwood <strong>forest</strong> species Ulmus minor due to the elm 

disease. Though this can not be confirmed for the surveyed area because <strong>of</strong> lacking data, 

a similar development may be assumed. 


43


1 0 0 m 

4 m 

Fig. 5.8 Cross section <strong>of</strong> the riverbed (lower fan) 

Ulmus minor regularly appeared, but never with higher importance. Large individuals 

were nearly completely lacking <strong>and</strong> only small juvenile trees were found. Occasionally 

larger stubs <strong>of</strong> Quercus robur were found, suggesting that this tree species due to its 

timber qualities is preferably logged <strong>and</strong> might have played a somewhat greater role 

within these <strong>forest</strong> types in the past. The Oplismenus undulatifolius-Acer velutinum 

community is similar in structure <strong>and</strong> thus related to a considerable degree with the 

Fraxinus excesior-Pterocarya pterocarpa community as demonstrated by the DCA <strong>and</strong> 

the NMDS (Fig. 4.23 <strong>and</strong> Fig. 4.24). 

The character species Acer velutinum is a large growing tree, that generally favours 

loamy <strong>and</strong> permanently moist soil conditions (Prilipko 1954). It is described to 

constitute <strong>forest</strong> st<strong>and</strong>s in river valleys in the Talysh-Lenkoran Zone (BIOFOR 2000). 

Few more information referring the ecology <strong>of</strong> this species is available. The closely 

related species Acer pseudoplatanus is known as a characteristic tree <strong>of</strong> scarcely flooded 

European lowl<strong>and</strong> <strong>forest</strong>s rich in nutrients (Späth 2007). Glaeser <strong>and</strong> Schmidt (2007) 

mention the strong dispersion <strong>of</strong> Acer pseudoplatanus <strong>and</strong> Fraxinus excelsior over large 

areas <strong>of</strong> the floodplain <strong>forest</strong> <strong>of</strong> Leipzig after dramatic hydrological modifications at the 

end <strong>of</strong> the 19 th century <strong>and</strong> emphasize the decreased importance <strong>of</strong> Quercus robur <strong>and</strong> 


44


Ulmus minor. One might suppose a similar development in the study area. Also 

remarkable is the scarcity <strong>of</strong> shrubs <strong>and</strong> trees <strong>of</strong> Salix spp. in the whole study area. This 

might be due to intensive grazing in <strong>and</strong> along the riverbeds but also to suppressed river 

dynamics <strong>and</strong> thus lacking rejuvenation options. 

5.2 Successional trends 

Based on the NMDS scatterplot (Fig. 4.24) a scheme was developed to show the 

successional trends in a simplified manner (Fig. 5.9). Successional development is 

represented by the second axis <strong>of</strong> the NMDS. The first axis mainly represents a gradient 

in constant soil moisture, which was also demonstrated by both DCA (Fig.4.22 <strong>and</strong> Fig. 

4.23). 

 

S u c c e s s i o n 

Q u e r c u s i b e r i c a - 

C a r p i n u s o r i e n t a l i s 

Q u e r c u s r o b u r - 

C a r p i n u s b e t u l u s 

O p l i s m e n u s u n d u l a t i f o l i u s - 

A c e r v e l u t i n u m 

F r a x i n u s e x c e l s i o r - 

P t e r o c a r y a p t e r o c a r p a 

 

S c l e r o p o a r i g i d a - 

P o p u l u s n i g r a 

E q u i s e t u m t e l m a t e i a - 

A l n u s b a r b a t a 

S w i d a s a n g u i n e a - 

P o p u l u s a l b a 

M 

o i s t u r e 

Fig. 5.9 Assumed successional development <strong>of</strong> the <strong>forest</strong> <strong>communities</strong> 

The lack <strong>of</strong> both bare soils <strong>and</strong> frequent floodings is responsible for the replacement <strong>of</strong> 

Scleropoa rigida-Populus nigra st<strong>and</strong>s by more competitive species regarding drought 

tolerance <strong>and</strong> germination conditions. Less flood tolerant species like Cotinus 


45

-Discussioncoggygria, 

Pyracantha coccinea, Crataegus spp. break the ground <strong>and</strong> subsequently 

tree species like Quercus iberica <strong>and</strong> Carpinus orientalis follow. Better water supplied 

<strong>and</strong> more nutrient rich site conditions might favour further succession to the Quercus 

robur-Carpinus betulus <strong>forest</strong>. Forest st<strong>and</strong>s <strong>of</strong> Quercus iberica-Carpinus orientalis are 

not subjected to successional trends. Rejuvenation on these sites quite well reflects the 

tree composition <strong>of</strong> the canopy plots. One might suggest that they represent a kind <strong>of</strong> 

terminal stage <strong>of</strong> <strong>forest</strong> development. Human activities like fuel wood cut <strong>and</strong> grazing 

might contribute to a certain part to this trend. The species composition <strong>of</strong> the Swida 

sanguinea-Populus alba rejuvenation differs completely from that one <strong>of</strong> its canopy 

plots. Succession leads to the Fraxinus excelsior-Pterocarya pterocarpa community. 

High rejuvenation rates <strong>of</strong> species like Carpinus betulus, Pterocarya pterocarpa <strong>and</strong> 

Crataegus spp. seem to be mainly responsible for this shift. In the beginning, river 

dynamics provide favourable conditions for Populus alba rejuvenation, mainly bare <strong>and</strong> 

permanently moist soils. On a once established Populus alba st<strong>and</strong> rejuvenation 

conditions for this species will worsen dramatically <strong>and</strong> more competitive species 

particularly regarding shade tolerance will germinate <strong>and</strong> consequently out compete all 

Populus alba, which are not fast enough to grow into the light <strong>and</strong> out <strong>of</strong> their reach. 

Finally, few huge Populus alba individuals will be left. The only chance for 

rejuvenation will be then by the way <strong>of</strong> root suckering, trying to find a recently 

originated gap in the tree layer. High rejuvenation rates <strong>of</strong> Swida sanguinea, Carpinus 

betulus, Fraxinus excelsior <strong>and</strong> Crataegus spp. suggest a possible succession <strong>of</strong> the 

Equisetum telmateia-Alnus barbata <strong>forest</strong> towards the Fraxinus excelsior-Pterocarya 

pterocarpa <strong>forest</strong>. But due to the <strong>of</strong>ten waterlogged conditions long term survival <strong>of</strong> 

those species is doubtful. Rejuvenation <strong>of</strong> Alnus barbata on these sites is scarce, since 

germination <strong>of</strong> this species is bound to bright light conditions (Brunner 2005), which 

are mainly lacking under closed canopy conditions. Regarding their tree species 

composition Fraxinus excelsior-Pterocarya pterocarpa <strong>and</strong> Oplismenus undulatifolius- 

Acer velutinum community are closely related. This is also reflected by their 

rejuvenation, which only slightly differs among themselves <strong>and</strong> from the respective 

canopy plots. Under the present conditions these <strong>forest</strong>s seem to constitute a kind <strong>of</strong> 

terminal stage <strong>of</strong> <strong>forest</strong> development on the well water <strong>and</strong> nutrient supplied soils <strong>of</strong> the 

lower fan. A number <strong>of</strong> parameters might contribute to this development, for example 

grazing or human impact in terms <strong>of</strong> hydrological changes, resulting in water table drop. 

Due to high human impact in terms <strong>of</strong> logging <strong>and</strong> grazing livestock rejuvenation within 


46

-Discussionthe 

Quercus robur-Carpinus betulus community is scarce. Juvenile Quercus robur 

individuals are completely lacking. Relatively high rejuvenation rates <strong>of</strong> rarely sampled 

species like Acer campestre, Mespilus germanica <strong>and</strong> Prunus cerasus might be 

responsible for a slight removal away from all other <strong>communities</strong> (Fig. 4.24 <strong>and</strong> 5.9). 

After all it should be kept in mind, that due to mainly small sized rejuvenation (<strong>of</strong>ten 

< 50 cm), the singularity <strong>of</strong> measurements <strong>and</strong> the relatively small dataset a serious 

prediction <strong>of</strong> <strong>forest</strong> succession can not be made. 

5.3 The human impact on the ecosystem 

A number <strong>of</strong> human activities take place within the study area, each with a more or less 

distinctive impact on the fan ecosystem. 

Flood protection 

Due to a number <strong>of</strong> flood protection measures, river dynamics on the alluvial fan near 

Qakh have been already considerably suppressed. Concrete walls <strong>of</strong> 2.5 metres height, 

gravel embankments <strong>and</strong> deepened channels (Fig. 5.10) restrict a great portion <strong>of</strong> the 

water discharge to the main-channel, thus preventing to a high extent channel 

Fig. 5.10 Concrete embankment (left) <strong>and</strong> channel deepening (right) 

instabilities (avulsions), which are <strong>of</strong> great importance for the maintenance <strong>of</strong> a cyclical 

succession regime (Gilvear et al. 2000). Annually in spring following the main flood 


47

-Discussionevent 

the government organizes reconstruction works at the main channel. Dozens <strong>of</strong> 

bulldozers are used to restore a stable <strong>and</strong> assessable discharge situation. 

Gravel dredging 

Dredging <strong>of</strong> gravels is intensively practised in the upper <strong>and</strong> mid-fan area. It is mainly 

used for road <strong>and</strong> building construction. Two plants, which are in private property 

already exist for about five years (Fig. 5.11). Another one was already working in 

former Soviet times <strong>and</strong> has recently been activated again. According to the chief 

workers about 200 m³ <strong>of</strong> gravel is daily mined mainly with old Soviet machinery, which 

has been acquired on very good terms. Due to the country's economic boom in the last 

decade there is actually an increasing dem<strong>and</strong> for gravel production. 

Fig. 5.11 Gravel dredging with old Russian machinery 

Tree logging <strong>and</strong> fuel wood cut 

The <strong>forest</strong>s <strong>of</strong> Azerbaijan are state property <strong>and</strong> due to their protective functions, 

designated as “<strong>forest</strong>s <strong>of</strong> the first group”. Hence, commercial logging is prohibited. The 

<strong>of</strong>ficially recorded (UN ECE. 2003) average annual <strong>forest</strong> cut was about 60,000 m³ for 

sanitary <strong>and</strong> maintenance purposes only. It was also estimated, that an additional 30,000 

to 40,000 m³ per year is cut illegally. The government <strong>of</strong> Azerbaijan has issued various 

regulations like the Forest Management Regulation, Af<strong>forest</strong>ation Regulation, 

Production Regulation <strong>and</strong> National Parks Regulation, but they contain many 

contradictions, ambiguities <strong>and</strong> gaps. As a result, they are little effective (WWF 2005). 


48


In all parts <strong>of</strong> the fan logging was observed. Populus alba <strong>and</strong> Quercus spp. are the 

most frequently logged trees (Fig. 5.12). Populus alba provides huge timber quantities 

<strong>of</strong> a relatively low quality, suitable mainly for ro<strong>of</strong> <strong>and</strong> floor construction. Quercus spp. 

provides less, but high quality wood for various uses. Other species, such as Acer 

velutinum, Alnus barbata, Carpinus betulus, Fraxinus excelsior <strong>and</strong> Pterocarya 

pterocarpa are also logged, but in much less quantity. A number <strong>of</strong> private sawmills 

around Qakh buy, cut <strong>and</strong> manufacture the timber brought by their clients. The 

provincial <strong>forest</strong>er in Qakh mentioned the sale <strong>of</strong> timber <strong>and</strong> estimated that about 6,000 

m³ <strong>of</strong> timber were sold in 2007. Timber prices are calculated on the basis <strong>of</strong> the former 

Russian system. He explained, that legally logged trees are marked before with a 

number given from the Forest Department. He also mentioned the occurrence <strong>of</strong> illegal 

logging mainly by night <strong>and</strong> the difficulties to prevent illegal logging. During our field 

work tree stubs with <strong>and</strong> also without a number were frequently found. People with 

horse <strong>and</strong> a cart full <strong>of</strong> fuel wood, which they had cut in the <strong>forest</strong> were met regularly 

(Fig. 5.12). Following independence <strong>and</strong> the collapse <strong>of</strong> the energy supply system, rural 

households became dependent on fuel wood for heating <strong>and</strong> cooking. According to 

statements <strong>of</strong> various people an annually consumption <strong>of</strong> at least 10 m³ <strong>of</strong> fuel wood per 

family must be assumed. Social <strong>and</strong> economic needs force people to illegally use the 

<strong>forest</strong> resources. Hence, impacts on <strong>forest</strong>s can not be taken seriously in an 

environmentally responsible way (WWF 2005). 

Fig. 5.12 Recently logged Populus alba (left) <strong>and</strong> cart filled with fuel wood (right) 


49


Wood pasture 

Grazing within the <strong>forest</strong> is common, but illegal. Despite existing financial penalties, we 

regularly observed herdsmen driving livestock through the <strong>forest</strong> <strong>and</strong> the riverbeds. On 

the upper fan it is mainly sheep, on the lower fan mostly cattle <strong>and</strong> water buffaloes (Fig. 

5.13). Some <strong>forest</strong> st<strong>and</strong>s are strongly affected by pasture, demonstrated by a very open 

structure <strong>and</strong> nearly lacking species recruitment. Particularly on the upper fan extensive 

areas are strongly grazed by sheep. The only wooden plants left are thorny shrubs like 

Paliurus spina christi, Punica granata <strong>and</strong> few Celtis caucasica provding shade for the 

livestock. The former vegetation was possibly constituted <strong>of</strong> Populus nigra <strong>and</strong> 

Quercus iberica-Carpinus orientalis <strong>forest</strong>. 

Since the enforcement <strong>of</strong> <strong>forest</strong> law seems to be ineffective <strong>and</strong> punishments are hardly 

executed, people will not ab<strong>and</strong>on wood pasture. 

Fig. 5.13 Water buffaloes in the poplar <strong>forest</strong> (right) <strong>and</strong> cattle in the riverbed (right) 

Plantations <strong>and</strong> Agriculture 

Soils <strong>of</strong> the lower fan are well water supplied <strong>and</strong> rich in nutrients. They are 

preferentially used for agriculture (see chapter 2.5). Apart from wheat, corn, melon <strong>and</strong> 

vegetable fields, plantations <strong>of</strong> hazelnut <strong>and</strong> walnut are abundant (Fig. 5.14). Hazelnut 

plantations are mainly located next to settlements <strong>and</strong> partly stretch out over extensive 

areas. Walnut plantations are <strong>of</strong>ten cultivated in the <strong>forest</strong> replacing the natural 

vegetation. We detected ab<strong>and</strong>oned or still cultivated plantations several times. They are 

administered by the Forest Department. Harvesting nuts in late summer <strong>and</strong> autumn 

provides a minimal income for rural people living in the surrounding villages. 


50


Fig. 5.14 Walnut plantation (left) <strong>and</strong> hazelnut plantation (right) 

5.4 Outlook 

Due to damaging activities like canalization, construction <strong>of</strong> flood embankments, 

agriculture <strong>and</strong> woodl<strong>and</strong> removal alluvial <strong>forest</strong> biodiversity has disappeared rapidly 

from the temperate regions <strong>of</strong> the world (Petts & Moller 1989; Wenger et al. 1990; 

Mitsch 1995). In Europe for example, the majority <strong>of</strong> alluvial <strong>forest</strong>s have long since 

been destroyed <strong>and</strong> many <strong>of</strong> the survivors are threatened by river regulation, power 

generation, pollution <strong>and</strong> intensified agriculture (Wenger et al. 1990). But these areas 

are particularly valuable ecosystems. They are characterized by a high biodiversity <strong>and</strong> 

sometimes high species richness within their habitat types (Large et al. 1994). Flood 

disturbance creates a variety <strong>of</strong> l<strong>and</strong> forms, which in turn allow a variety <strong>of</strong> woodl<strong>and</strong> 

types to co-exist with mosaics <strong>of</strong> open vegetation. The disturbance itself helps to 

maintain a range <strong>of</strong> successional stages. Numerous ecotones persist between both, 

various types <strong>of</strong> terrestrial vegetation <strong>and</strong> aquatic habitats. Interactions between l<strong>and</strong> 

<strong>and</strong> water permanently occur in a variety <strong>of</strong> ways, such as the input <strong>of</strong> dead wood or 

other organic matter into aquatic habitats. The constantly high level <strong>of</strong> soil moisture 

maintains high biological productivity. Riparian zones form corridors for wildlife 

movement <strong>and</strong> thus important links in the l<strong>and</strong>scape (Peterken & Hughes 1995). 

The conservation <strong>of</strong> these areas requires sympathetic environmental management <strong>and</strong> 

must not only examine l<strong>and</strong> use <strong>and</strong> activity within the river corridor, but also consider 

impacts on flow regime, flood magnitude <strong>and</strong> frequency, at the catchment scale (Gilvear 

et al. 2000). 


51

-Summary- 

6 Summary 

With this study structure, species composition <strong>and</strong> successional trends <strong>of</strong> alluvial <strong>forest</strong> 

<strong>communities</strong> in northwestern Azerbaijan were analysed. Research object was an alluvial 

fan situated at the southern slope <strong>of</strong> the Greater Caucasus. Compared to other fans <strong>of</strong> 

this region a considerable amount <strong>of</strong> <strong>forest</strong>s is still preserved. However, despite existing 

penalties wood pasture <strong>and</strong> wood logging are common. Based on both 71 vegetation 

samples <strong>and</strong> 59 samples <strong>of</strong> <strong>forest</strong> structure eight plant <strong>communities</strong> were distinguished 

by means <strong>of</strong> a cluster analysis: Equisetum telmateia-Alnus barbata community, 

Fraxinus excelsior-Pterocarya pterocarpa com., Swida sanguinea-Populus alba com., 

Oplismenus undulatifolius-Acer velutinum com., Quercus robur-Carpinus betulus com., 

Quercus iberica-Carpinus-orientalis com., Scleropoa rigida-Populus nigra com., 

Echinochloa crusgalli-Populus alba com. By means <strong>of</strong> indirect ordination (DCA) 

location on the fan, water supply <strong>and</strong> soil texture were revealed as the most important 

factors for the establishment <strong>of</strong> the different vegetation types. St<strong>and</strong>s <strong>of</strong> Populus nigra 

<strong>and</strong> Populus alba are restricted to frequently flooded sites next to the riverbed. The 

former one is more drought resistant <strong>and</strong> thus higher located than the latter one. Their 

<strong>regeneration</strong> strongly depends on the presence <strong>of</strong> bare soils created by natural river 

dynamics. The thermophile Quercus iberica-Carpinus orientalis community is the least 

affected by flooding <strong>and</strong> replaces Populus nigra st<strong>and</strong>s in case <strong>of</strong> disappearing river 

dynamics. The Quercus robur-Carpinus betulus community is mainly restricted to the 

the mid-fan. Strong human impact is responsible for nearly lacking <strong>regeneration</strong>. The 

presence <strong>of</strong> Quercus robur in lower fan sections is low. But due to a preferential 

logging <strong>of</strong> oak it might have been considerably higher in the past. Alnus barbata st<strong>and</strong>s 

are mainly restricted to the mid-fan, where permanent springs cause constantly 

waterlogged soil conditions. The lower fan is mainly covered with mixed st<strong>and</strong>s <strong>of</strong> Acer 

velutinum, Carpinus betulus, Fraxinus excelsior <strong>and</strong> Pterocarya pterocarpa, which, 

according to the rejuvenation at those sites, will also dominate in the future. At the very 

same sites a few huge Populus alba individuals are remnants <strong>of</strong> former open riverbed 

<strong>communities</strong>, which are to rejuvenate in closed canopy st<strong>and</strong>s. Due to its high rate <strong>of</strong> 

growth <strong>and</strong> enormous timber volume it is apart from oak the most frequently logged 

tree species on the fan. 

It has been shown that flood protection measures, gravel dredging, logging, fuel wood 

cut, pasture <strong>and</strong> agriculture have a more or less distinctive negative impact on the 


52

-Summaryecosystem. 

Hence, intensive efforts have to be made to preserve this highly dynamic 

ecosystem. It provides a mosaic <strong>of</strong> many different habitats, which is diverse in itself <strong>and</strong> 

different from the <strong>forest</strong>s <strong>of</strong> adjacent ecosystems. 

Fig. 6.1 River dynamics on the upper fan with Populus nigra in the front 


53

-Zusammenfassung- 

7 Zusammenfassung 

Im nordwestlichen Teil von Aserbaidschan am Südabhang des Großen Kaukasus 

wurden Waldgesellschaften auf einem vergleichsweise gering genutzten 

Schwemmfächer hinsichtlich ihrer Artenzusammensetzung, Best<strong>and</strong>esstruktur und 

Verjüngung untersucht. Basierend auf 71 Vegetationsaufnahmen und 59 Aufnahmen 

zur Best<strong>and</strong>esstruktur wurden die folgenden acht Pflanzengesellschaften mittels 

Clusteranalyse ausgeschieden: Equisetum telmateia-Alnus barbata-Gesellschaft, 

Fraxinus excelsior-Pterocarya pterocarpa-Gesellschaft, Swida sanguinea-Populus 

alba-Gesellschaft, Oplismenus undulatifolius-Acer velutinum-Gesellschaft, Quercus 

robur-Carpinus betulus-Gesellschaft, Quercus iberica-Carpinus orientalis-Gesellschaft, 

Scleropoa rigida-Populus nigra-Gesellschaft und Echinochloa crusgalli-Populus alba- 

Gesellschaft. Die Einflüsse der abiotischen St<strong>and</strong>ortfaktoren wurden durch indirekte 

Ordination (DCA) ermittelt. Als wichtigste prägende Faktoren wurden Lage auf dem 

Fächer, Substratbeschaffenheit und Bodenfeuchte ermittelt. Sowohl Populus nigra- als 

auch Populus alba-Gesellschaft sind auf die unmittelbare Nähe zum Flusslauf 

angewiesen, wobei erstere höher auf dem Schwemmfächer gelegene, wechselfeuchte 

und letztere niedriger gelegene, permanent feuchte St<strong>and</strong>orte besiedelt. Um sich zu 

verjüngen, sind beide Gesellschaften auf die natürliche Flußdynamik und durch sie 

geschaffene Rohböden angewiesen. Die wärmeliebende Quercus iberica-Carpinus 

orientalis-Gesellschaft besiedelt ausschließlich St<strong>and</strong>orte im oberen 

Schwemmfächerabschnitt. Sie bildet die Schlußgesellschaft auf höher gelegenen, 

grobsubstratreichen, von Überflutungen wenig beeinflußten St<strong>and</strong>orten und ist 

Sukzessionsfolger auf nicht mehr von der Flußdynamik „gestörten“ Populus nigra 

St<strong>and</strong>orten. Auf feineren und frischeren Böden wird sie von der Quercus robur- 

Carpinus betulus-Gesellschaft abgelöst. Starke Beweidung und hoher menschlicher 

Nutzungsdruck sind hier verantwortlich für das fast vollständige Fehlen von 

Verjüngung. Möglicherweise kam Quercus robur früher weitaus häufiger auch im 

unteren Schwemmfächerabschnitt vor. Vereinzelt gefundene, groß gewachsene 

Exemplare sowie die für das gesamte Gebiet geltende starke Nutzung von Eichen 

könnten Indizien dafür sein. Das Vorkommen der Equisetum telmateia-Alnus barbata- 

Gesellschaft ist vorwiegend auf den mittleren Fächerabschnitt beschränkt, wo 

Grundwasser an zahlreichen permanenten Quellen austritt. Im unteren Fächerabschnitt 


54

-Zusammenfassungdominieren 

gemischte Bestände von Acer velutinum, Carpinus betulus, Fraxinus 

excelsior und Pterocarya pterocarpa, die sich auch auch in der Verjüngung als 

dominant erweisen. Wenige, hochgewachsene und <strong>and</strong>ere Arten weit überragende 

Populus alba-Exemplare zeugen von ehemals <strong>of</strong>fenen, der Flussdynamik unterworfenen 

St<strong>and</strong>orten. Mit zunehmender Bewaldung dieser St<strong>and</strong>orte unterbleibt jedoch die 

Verjüngung dieser Art, sie ist auf längere Sicht nicht mehr konkurrenzfähig. Aufgrund 

ihrer hohen Zuwachsraten und riesigen Holzvolumen ist Populus alba ebenfalls massiv 

vom Holzeinschlag betr<strong>of</strong>fen. 

Die natürlichen Abläufe dieses hochdynamischen Ökosystems werden durch 

Maßnahmen zur Überflutungssicherung, Holzeinschlag, Schotterabbau, Beweidung und 

l<strong>and</strong>wirtschaftliche Nutzung beeinträchtigt. Es müssen daher Mittel und Wege gefunden 

werden, diese Funktionen auch in Zukunft im Einklang mit verantwortlicher 

menschlicher Nutzung zu garantieren. 

Fig. 7.1 Natural riverbed with Populus alba 


55

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59

-Appendix I - 

Appendix I 

Structure parameters <strong>of</strong> the <strong>forest</strong> <strong>communities</strong> 

1. Equisetum telmateia-Alnus barbata community 

Species Total ALN-BAR CAR-BET CAR-ORI CRA-SPP DIO-LOT FIC-CAR FRA-EXC JUG-REG MOR-SPP POP-ALB PTE-PTE SAL-SPP SWI-SAN ULM-MIN 

Number 220 157 4 8 8 3 1 5 2 3 7 7 2 11 2 

Mean Distance [m] 5 / / / / / / / / / / / / / / 

Mean DBH [cm] / 36 23 16 9 6 8 22 29 10 97 30 19 6 17 

Portion <strong>of</strong> Stock [%] / 71 2 4 4 1 0 2 1 1 3 3 1 5 1 

Number/ha 548 426 11 23 18 5 1 11 2 3 10 9 5 21 4 

Mean (B)asal (A)rea [cm²] / 1041 412 193 60 32 50 393 641 86 7391 693 291 31 216 

Total BA [m²/ha] 46 36 0 0 0 0 0 0 0 0 7 1 0 0 0 

Portion <strong>of</strong> Total BA/ha [%] / 78 1 1 0 0 0 1 0 0 16 2 0 0 0 

Absolute Frequency [%] 178 91 5 5 9 5 2 7 4 4 9 9 4 20 4 

Relative Frequency [%] / 51 3 3 5 3 1 4 2 2 5 5 2 11 2 

Importance Value / 213 6 8 7 4 1 6 3 3 19 10 3 15 3 

2. Fraxinus excelsior-Pterocarya pterocarpa community 

Species Total ACE-CAM ACE-VEL ALN-BAR CAR-BET CAR-ORI COR-AVE CRA-SPP EUO-EUR FRA-EXC MES-GER POP-ALB QUE-ROB PRU-CER PTE-PTE SWI-SAN ULM-MIN 

Number 220 2 26 17 59 1 13 8 2 42 1 5 1 4 25 10 4 

Mean Distance [m] 5 / / / / / / / / / / / / 

Mean DBH [cm] / 4 28 35 35 10 12 18 4 34 18 103 37 12 38 6 20 

Portion <strong>of</strong> Stock [%] / 1 12 8 27 0 6 4 1 19 0 2 0 2 11 5 2 

Number/ha 482 3 59 45 116 1 31 12 8 91 1 5 1 8 65 31 6 

Mean (B)asal (A)rea [cm²] / 12 603 977 963 81 117 258 14 888 249 8267 1071 104 1141 24 309 

Total BA [m²/ha] 37 0 4 4 10 0 0 0 0 6 0 4 0 0 7 0 0 

Portion <strong>of</strong> Total BA/ha [%] / 0 11 12 26 0 1 1 0 17 0 12 0 0 18 0 0 

Absolute Frequency [%] 269 4 35 22 55 2 22 11 4 42 2 5 2 7 38 15 5 

Relative Frequency [%] / 1 13 8 20 1 8 4 1 16 1 2 1 3 14 5 2 

Importance Value / 3 34 26 76 1 14 10 2 57 1 16 2 5 41 9 4 

3. Swida sanguinea-Populus alba community 

Species 

Total ACE-VEL ALN-BAR CAR-ORI COR-AVE CRA-SPP DIO-LOT FIC-CAR JUG-REG MOR-SPP POP-ALB POP-NIG PRU-CER PTE-PTE SAL-SPP SWI-SAN ULM-MIN 

Number 180 2 5 3 2 2 1 2 2 21 84 11 5 22 2 10 6 


Mean DBH [cm] / 13 27 7 8 7 5 15 5 9 37 14 6 15 21 7 11 

Portion <strong>of</strong> Stock [%] / 1 3 2 1 1 1 1 1 12 47 6 3 12 1 6 3 

Number/ha 792 7 9 24 7 18 3 3 3 55 446 75 18 59 9 30 25 

Mean (B)asal (A)rea [cm²] / 143 557 41 47 44 23 170 22 67 1094 151 25 182 331 38 92 

Total BA [m²/ha] 36 0 0 0 0 0 0 0 0 0 32 1 0 1 0 0 0 

Portion <strong>of</strong> Total BA/ha [%] / 0 1 0 0 0 0 0 0 1 90 3 0 3 0 0 1 

Absolute Frequency [%] 231 4 7 7 4 4 2 2 4 27 78 16 11 29 4 18 13 

Relative Frequency [%] / 2 3 3 2 2 1 1 2 12 34 7 5 13 2 8 6 

Importance Value / 3 7 5 4 3 1 2 3 24 172 16 7 27 3 13 11 

4. Oplismenus undulatifolius-Acer velutinum community 

Species 

Total ACE-CAM ULM-MIN FRA-EXC SWI-SAN ACE-VEL COR-AVE PTE-PTE PRU-CER CAR-BET QUE-ROB 

Number 80 2 4 8 6 33 5 4 1 16 1 

Mean Distance [m] 5 / / / / / / / / / / 

Mean DBH [cm] / 5 6 44 4 41 8 26 6 38 102 

Portion <strong>of</strong> Stock [%] / 3 5 10 8 41 6 5 1 20 1 

Number/ha 541 16 27 50 57 212 38 37 11 88 3 

Mean (B)asal (A)rea [cm²] / 17 29 1500 16 1299 49 548 29 1116 8251 

Total BA [m²/ha] 46 0 0 6 0 26 0 1 0 11 3 

Portion <strong>of</strong> Total BA/ha [%] / 0 0 12 0 56 0 2 0 24 6 

Absolute Frequency [%] 260 10 15 30 25 75 20 15 5 60 5 

Relative Frequency [%] / 4 6 12 10 29 8 6 2 23 2 

Importance Value / 6 11 33 17 128 13 13 3 66 9 

5. Quercus robur-Carpinus betulus community 

Species 

Total ACE-CAM CAR-BET CAR-ORI COR-AVE COR-MAS CRA-SPP CYD-OBL FRA-EXC MES-GER MOR-SPP POP-ALB PRU-CER QUE-IBE QUE-ROB SWI-SAN 

Number 100 3 40 10 4 1 2 1 2 5 1 2 1 4 18 6 

Mean Distance [m] 5 / / / / / / / / / / / / / / / 

Mean DBH [cm] / 20 35 19 11 6 35 14 10 12 7 79 4 44 85 8 

Portion <strong>of</strong> Stock [%] / 3 40 10 4 1 2 1 2 5 1 2 1 4 18 6 

Number/ha 362 11 139 32 17 3 6 3 7 19 5 8 5 13 69 25 

Mean (B)asal (A)rea [cm²] / 320 954 277 99 32 966 161 84 107 42 4863 11 1521 5639 51 

Total BA [m²/ha] 61 0 13 1 0 0 1 0 0 0 0 4 0 2 40 0 

Portion <strong>of</strong> Total BA/ha [%] / 1 21 1 0 0 1 0 0 0 0 6 0 3 66 0 

Absolute Frequency [%] 268 12 72 24 16 4 4 4 8 20 4 8 4 12 56 20 

Relative Frequency [%] / 4 27 9 6 1 1 1 3 7 1 3 1 4 21 7 

Importance Value / 9 95 22 10 2 5 2 5 12 2 12 2 15 93 13 


60

-Appendix I - 

6. Quercus iberica-Carpinus orientalis community 

Species 

Total ACE-CAM CAR-BET CAR-ORI COR-MAS COT-COG CRA-SPP MES-GER POP-ALB POP-NIG QUE-IBE QUE-ROB ULM-MIN 

Number 320 4 1 178 1 23 38 1 3 5 56 2 8 


Mean DBH [cm] / 20 21 13 4 8 10 7 63 46 32 32 16 

Portion <strong>of</strong> Stock [%] / 1 0 56 0 7 12 0 1 2 18 1 3 

Number/ha 979 10 3 581 1 70 104 4 6 12 163 6 20 

Mean (B)asal (A)rea [cm²] / 301 336 140 11 48 82 42 3145 1669 811 784 189 

Total BA [m²/ha] 25 0 0 7 0 0 1 0 2 1 12 0 0 

Portion <strong>of</strong> Total BA/ha [%] / 1 0 29 0 1 4 0 6 5 49 2 2 

Absolute Frequency [%] 215 4 1 91 1 18 30 1 4 6 46 3 10 

Relative Frequency [%] / 2 1 42 1 8 14 1 2 3 22 1 5 

Importance Value / 4 1 134 1 21 29 1 6 8 85 3 8 

7. Scleropoa rigida-Populus nigra community 

Species 

Total CAR-ORI COT-COG CRA-SPP HIP-RHA LIG-VUL MES-GER POP-ALB POP-NIG PYR-COC QUE-IBE ROB-PSE 

Number 60 2 21 2 5 3 1 2 17 1 3 3 

Mean Distance [m] 4 / / / / / / / / / / / 

Mean DBH [cm] / 8 6 4 6 4 7 37 24 6 19 12 

Portion <strong>of</strong> Stock [%] / 3 35 3 8 5 2 3 28 2 5 5 

Number/ha 692 17 270 17 61 52 9 18 172 8 34 34 

Mean (B)asal (A)rea [cm²] / 52 30 16 30 14 36 1080 437 29 280 114 

Total BA [m²/ha] 11 0 1 0 0 0 0 2 7 0 1 0 

Portion <strong>of</strong> Total BA/ha [%] / 1 7 0 1 1 0 17 62 0 6 3 

Absolute Frequency [%] 273 13 73 13 33 7 7 13 73 7 20 13 

Relative Frequency [%] / 5 27 5 12 2 2 5 27 2 7 5 

Importance Value / 9 72 8 23 9 4 23 116 4 18 14 


61

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